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WAPL_DROME
ID   WAPL_DROME              Reviewed;        1741 AA.
AC   Q9W517; O46095; Q24153; Q7KVZ8; Q8MRI2; Q9W518;
DT   06-DEC-2005, integrated into UniProtKB/Swiss-Prot.
DT   01-MAY-2000, sequence version 1.
DT   03-AUG-2022, entry version 137.
DE   RecName: Full=Protein wings apart-like;
DE   AltName: Full=Protein parallel sister chromatids;
GN   Name=wapl {ECO:0000312|EMBL:AAF45734.1};
GN   Synonyms=pasc {ECO:0000303|PubMed:10747063}; ORFNames=CG3707;
OS   Drosophila melanogaster (Fruit fly).
OC   Eukaryota; Metazoa; Ecdysozoa; Arthropoda; Hexapoda; Insecta; Pterygota;
OC   Neoptera; Endopterygota; Diptera; Brachycera; Muscomorpha; Ephydroidea;
OC   Drosophilidae; Drosophila; Sophophora.
OX   NCBI_TaxID=7227;
RN   [1] {ECO:0000305, ECO:0000312|EMBL:AAA91230.1}
RP   NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM A), FUNCTION, DEVELOPMENTAL STAGE, AND
RP   DISRUPTION PHENOTYPE.
RC   TISSUE=Embryo {ECO:0000269|PubMed:10747063}, and Imaginal disk;
RX   PubMed=10747063; DOI=10.1093/genetics/154.4.1693;
RA   Verni F., Gandhi R., Goldberg M.L., Gatti M.;
RT   "Genetic and molecular analysis of wings apart-like (wapl), a gene
RT   controlling heterochromatin organization in Drosophila melanogaster.";
RL   Genetics 154:1693-1710(2000).
RN   [2] {ECO:0000312|EMBL:AAF45734.1}
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC   STRAIN=Berkeley {ECO:0000269|PubMed:10731132};
RX   PubMed=10731132; DOI=10.1126/science.287.5461.2185;
RA   Adams M.D., Celniker S.E., Holt R.A., Evans C.A., Gocayne J.D.,
RA   Amanatides P.G., Scherer S.E., Li P.W., Hoskins R.A., Galle R.F.,
RA   George R.A., Lewis S.E., Richards S., Ashburner M., Henderson S.N.,
RA   Sutton G.G., Wortman J.R., Yandell M.D., Zhang Q., Chen L.X., Brandon R.C.,
RA   Rogers Y.-H.C., Blazej R.G., Champe M., Pfeiffer B.D., Wan K.H., Doyle C.,
RA   Baxter E.G., Helt G., Nelson C.R., Miklos G.L.G., Abril J.F., Agbayani A.,
RA   An H.-J., Andrews-Pfannkoch C., Baldwin D., Ballew R.M., Basu A.,
RA   Baxendale J., Bayraktaroglu L., Beasley E.M., Beeson K.Y., Benos P.V.,
RA   Berman B.P., Bhandari D., Bolshakov S., Borkova D., Botchan M.R., Bouck J.,
RA   Brokstein P., Brottier P., Burtis K.C., Busam D.A., Butler H., Cadieu E.,
RA   Center A., Chandra I., Cherry J.M., Cawley S., Dahlke C., Davenport L.B.,
RA   Davies P., de Pablos B., Delcher A., Deng Z., Mays A.D., Dew I.,
RA   Dietz S.M., Dodson K., Doup L.E., Downes M., Dugan-Rocha S., Dunkov B.C.,
RA   Dunn P., Durbin K.J., Evangelista C.C., Ferraz C., Ferriera S.,
RA   Fleischmann W., Fosler C., Gabrielian A.E., Garg N.S., Gelbart W.M.,
RA   Glasser K., Glodek A., Gong F., Gorrell J.H., Gu Z., Guan P., Harris M.,
RA   Harris N.L., Harvey D.A., Heiman T.J., Hernandez J.R., Houck J., Hostin D.,
RA   Houston K.A., Howland T.J., Wei M.-H., Ibegwam C., Jalali M., Kalush F.,
RA   Karpen G.H., Ke Z., Kennison J.A., Ketchum K.A., Kimmel B.E., Kodira C.D.,
RA   Kraft C.L., Kravitz S., Kulp D., Lai Z., Lasko P., Lei Y., Levitsky A.A.,
RA   Li J.H., Li Z., Liang Y., Lin X., Liu X., Mattei B., McIntosh T.C.,
RA   McLeod M.P., McPherson D., Merkulov G., Milshina N.V., Mobarry C.,
RA   Morris J., Moshrefi A., Mount S.M., Moy M., Murphy B., Murphy L.,
RA   Muzny D.M., Nelson D.L., Nelson D.R., Nelson K.A., Nixon K., Nusskern D.R.,
RA   Pacleb J.M., Palazzolo M., Pittman G.S., Pan S., Pollard J., Puri V.,
RA   Reese M.G., Reinert K., Remington K., Saunders R.D.C., Scheeler F.,
RA   Shen H., Shue B.C., Siden-Kiamos I., Simpson M., Skupski M.P., Smith T.J.,
RA   Spier E., Spradling A.C., Stapleton M., Strong R., Sun E., Svirskas R.,
RA   Tector C., Turner R., Venter E., Wang A.H., Wang X., Wang Z.-Y.,
RA   Wassarman D.A., Weinstock G.M., Weissenbach J., Williams S.M., Woodage T.,
RA   Worley K.C., Wu D., Yang S., Yao Q.A., Ye J., Yeh R.-F., Zaveri J.S.,
RA   Zhan M., Zhang G., Zhao Q., Zheng L., Zheng X.H., Zhong F.N., Zhong W.,
RA   Zhou X., Zhu S.C., Zhu X., Smith H.O., Gibbs R.A., Myers E.W., Rubin G.M.,
RA   Venter J.C.;
RT   "The genome sequence of Drosophila melanogaster.";
RL   Science 287:2185-2195(2000).
RN   [3] {ECO:0000305, ECO:0000312|EMBL:AAF45734.1}
RP   GENOME REANNOTATION, AND ALTERNATIVE SPLICING.
RC   STRAIN=Berkeley;
RX   PubMed=12537572; DOI=10.1186/gb-2002-3-12-research0083;
RA   Misra S., Crosby M.A., Mungall C.J., Matthews B.B., Campbell K.S.,
RA   Hradecky P., Huang Y., Kaminker J.S., Millburn G.H., Prochnik S.E.,
RA   Smith C.D., Tupy J.L., Whitfield E.J., Bayraktaroglu L., Berman B.P.,
RA   Bettencourt B.R., Celniker S.E., de Grey A.D.N.J., Drysdale R.A.,
RA   Harris N.L., Richter J., Russo S., Schroeder A.J., Shu S.Q., Stapleton M.,
RA   Yamada C., Ashburner M., Gelbart W.M., Rubin G.M., Lewis S.E.;
RT   "Annotation of the Drosophila melanogaster euchromatic genome: a systematic
RT   review.";
RL   Genome Biol. 3:RESEARCH0083.1-RESEARCH0083.22(2002).
RN   [4] {ECO:0000305, ECO:0000312|EMBL:CAB10973.1}
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC   STRAIN=Oregon-R {ECO:0000269|PubMed:10731137};
RX   PubMed=10731137; DOI=10.1126/science.287.5461.2220;
RA   Benos P.V., Gatt M.K., Ashburner M., Murphy L., Harris D., Barrell B.G.,
RA   Ferraz C., Vidal S., Brun C., Demailles J., Cadieu E., Dreano S., Gloux S.,
RA   Lelaure V., Mottier S., Galibert F., Borkova D., Minana B., Kafatos F.C.,
RA   Louis C., Siden-Kiamos I., Bolshakov S., Papagiannakis G., Spanos L.,
RA   Cox S., Madueno E., de Pablos B., Modolell J., Peter A., Schoettler P.,
RA   Werner M., Mourkioti F., Beinert N., Dowe G., Schaefer U., Jaeckle H.,
RA   Bucheton A., Callister D.M., Campbell L.A., Darlamitsou A., Henderson N.S.,
RA   McMillan P.J., Salles C., Tait E.A., Valenti P., Saunders R.D.C.,
RA   Glover D.M.;
RT   "From sequence to chromosome: the tip of the X chromosome of D.
RT   melanogaster.";
RL   Science 287:2220-2222(2000).
RN   [5] {ECO:0000305, ECO:0000312|EMBL:AAM50260.1}
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM A).
RC   STRAIN=Berkeley {ECO:0000312|EMBL:AAM50260.1};
RC   TISSUE=Embryo {ECO:0000269|PubMed:12537569};
RX   PubMed=12537569; DOI=10.1186/gb-2002-3-12-research0080;
RA   Stapleton M., Carlson J.W., Brokstein P., Yu C., Champe M., George R.A.,
RA   Guarin H., Kronmiller B., Pacleb J.M., Park S., Wan K.H., Rubin G.M.,
RA   Celniker S.E.;
RT   "A Drosophila full-length cDNA resource.";
RL   Genome Biol. 3:RESEARCH0080.1-RESEARCH0080.8(2002).
RN   [6]
RP   PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT THR-258; SER-355 AND SER-888, AND
RP   IDENTIFICATION BY MASS SPECTROMETRY.
RC   TISSUE=Embryo;
RX   PubMed=18327897; DOI=10.1021/pr700696a;
RA   Zhai B., Villen J., Beausoleil S.A., Mintseris J., Gygi S.P.;
RT   "Phosphoproteome analysis of Drosophila melanogaster embryos.";
RL   J. Proteome Res. 7:1675-1682(2008).
CC   -!- FUNCTION: Has a role in female meiotic chromosome segregation in
CC       females; proximal heterochromatin is involved in chromosome pairing
CC       during female meiosis. Is a dominant suppressor of both white and
CC       Stubble position-effect variegation (PEV), while it is a weak enhancer
CC       of brown variegation. {ECO:0000269|PubMed:10747063}.
CC   -!- ALTERNATIVE PRODUCTS:
CC       Event=Alternative splicing; Named isoforms=2;
CC       Name=A {ECO:0000269|PubMed:10747063};
CC         IsoId=Q9W517-1; Sequence=Displayed;
CC       Name=B {ECO:0000303|PubMed:10731132};
CC         IsoId=Q9W517-2; Sequence=VSP_051920;
CC   -!- DEVELOPMENTAL STAGE: Expressed in embryos.
CC       {ECO:0000269|PubMed:10747063}.
CC   -!- DISRUPTION PHENOTYPE: Flies exhibit brain cell metaphases where sister
CC       chromatids of all chromosomes are aligned parallel to each other
CC       instead of assuming the typical morphology, heterochromatin
CC       condensation or chromosome segregation are not affected.
CC       {ECO:0000269|PubMed:10747063}.
CC   -!- SIMILARITY: Belongs to the WAPL family. {ECO:0000305}.
CC   -!- SEQUENCE CAUTION:
CC       Sequence=AAA91230.1; Type=Frameshift; Evidence={ECO:0000305};
CC       Sequence=AAM50260.1; Type=Frameshift; Evidence={ECO:0000305};
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DR   EMBL; U40214; AAA91230.1; ALT_FRAME; mRNA.
DR   EMBL; AE014298; AAF45734.1; -; Genomic_DNA.
DR   EMBL; AE014298; AAF45735.2; -; Genomic_DNA.
DR   EMBL; Z98269; CAB10973.1; -; Genomic_DNA.
DR   EMBL; AY119606; AAM50260.1; ALT_FRAME; mRNA.
DR   PIR; T13349; T13349.
DR   PIR; T13610; T13610.
DR   RefSeq; NP_001284804.1; NM_001297875.1. [Q9W517-1]
DR   RefSeq; NP_525042.2; NM_080303.3. [Q9W517-1]
DR   RefSeq; NP_726796.2; NM_166931.2. [Q9W517-2]
DR   AlphaFoldDB; Q9W517; -.
DR   SMR; Q9W517; -.
DR   BioGRID; 57728; 2.
DR   IntAct; Q9W517; 1.
DR   STRING; 7227.FBpp0070413; -.
DR   iPTMnet; Q9W517; -.
DR   PaxDb; Q9W517; -.
DR   EnsemblMetazoa; FBtr0070429; FBpp0070413; FBgn0004655. [Q9W517-1]
DR   EnsemblMetazoa; FBtr0070430; FBpp0070414; FBgn0004655. [Q9W517-2]
DR   EnsemblMetazoa; FBtr0344159; FBpp0310572; FBgn0004655. [Q9W517-1]
DR   GeneID; 31187; -.
DR   KEGG; dme:Dmel_CG3707; -.
DR   CTD; 23063; -.
DR   FlyBase; FBgn0004655; wapl.
DR   VEuPathDB; VectorBase:FBgn0004655; -.
DR   eggNOG; KOG2152; Eukaryota.
DR   GeneTree; ENSGT00390000015768; -.
DR   HOGENOM; CLU_001745_0_0_1; -.
DR   InParanoid; Q9W517; -.
DR   OMA; IQQLPHC; -.
DR   PhylomeDB; Q9W517; -.
DR   Reactome; R-DME-2468052; Establishment of Sister Chromatid Cohesion.
DR   Reactome; R-DME-2470946; Cohesin Loading onto Chromatin.
DR   Reactome; R-DME-2500257; Resolution of Sister Chromatid Cohesion.
DR   BioGRID-ORCS; 31187; 0 hits in 1 CRISPR screen.
DR   GenomeRNAi; 31187; -.
DR   PRO; PR:Q9W517; -.
DR   Proteomes; UP000000803; Chromosome X.
DR   Bgee; FBgn0004655; Expressed in cleaving embryo and 24 other tissues.
DR   ExpressionAtlas; Q9W517; baseline and differential.
DR   Genevisible; Q9W517; DM.
DR   GO; GO:0005634; C:nucleus; IDA:FlyBase.
DR   GO; GO:0005700; C:polytene chromosome; IDA:FlyBase.
DR   GO; GO:0006325; P:chromatin organization; IMP:UniProtKB.
DR   GO; GO:0051321; P:meiotic cell cycle; IEA:UniProtKB-KW.
DR   GO; GO:0061781; P:mitotic cohesin unloading; IMP:FlyBase.
DR   GO; GO:0060623; P:regulation of chromosome condensation; IBA:GO_Central.
DR   GO; GO:0071922; P:regulation of cohesin loading; IBA:GO_Central.
DR   GO; GO:0007062; P:sister chromatid cohesion; IMP:FlyBase.
DR   Gene3D; 1.25.10.10; -; 1.
DR   InterPro; IPR011989; ARM-like.
DR   InterPro; IPR016024; ARM-type_fold.
DR   InterPro; IPR039874; WAPL.
DR   InterPro; IPR022771; WAPL_C.
DR   InterPro; IPR012502; WAPL_dom.
DR   PANTHER; PTHR22100; PTHR22100; 2.
DR   Pfam; PF07814; WAPL; 1.
DR   SUPFAM; SSF48371; SSF48371; 1.
DR   PROSITE; PS51271; WAPL; 1.
PE   1: Evidence at protein level;
KW   Alternative splicing; Chromosome partition; Meiosis; Phosphoprotein;
KW   Reference proteome.
FT   CHAIN           1..1741
FT                   /note="Protein wings apart-like"
FT                   /id="PRO_0000076364"
FT   DOMAIN          1140..1648
FT                   /note="WAPL"
FT                   /evidence="ECO:0000255|PROSITE-ProRule:PRU00603"
FT   REGION          68..100
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   REGION          119..277
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   REGION          291..472
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   REGION          490..612
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   REGION          675..810
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   REGION          823..963
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   REGION          1038..1068
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   REGION          1112..1141
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   REGION          1708..1741
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        68..83
FT                   /note="Polar residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        217..233
FT                   /note="Basic and acidic residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        242..260
FT                   /note="Basic and acidic residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        309..333
FT                   /note="Polar residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        349..380
FT                   /note="Polar residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        439..456
FT                   /note="Acidic residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        530..612
FT                   /note="Polar residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        845..860
FT                   /note="Polar residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        871..898
FT                   /note="Basic and acidic residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        899..917
FT                   /note="Polar residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   COMPBIAS        1046..1068
FT                   /note="Polar residues"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   MOD_RES         258
FT                   /note="Phosphothreonine"
FT                   /evidence="ECO:0000269|PubMed:18327897"
FT   MOD_RES         355
FT                   /note="Phosphoserine"
FT                   /evidence="ECO:0000269|PubMed:18327897"
FT   MOD_RES         888
FT                   /note="Phosphoserine"
FT                   /evidence="ECO:0000269|PubMed:18327897"
FT   VAR_SEQ         1..649
FT                   /note="Missing (in isoform B)"
FT                   /evidence="ECO:0000303|PubMed:10731132"
FT                   /id="VSP_051920"
FT   CONFLICT        37
FT                   /note="W -> L (in Ref. 5; AAM50260)"
FT                   /evidence="ECO:0000305"
FT   CONFLICT        135
FT                   /note="A -> V (in Ref. 4; CAB10973)"
FT                   /evidence="ECO:0000305"
FT   CONFLICT        772
FT                   /note="T -> M (in Ref. 4; CAB10973)"
FT                   /evidence="ECO:0000305"
FT   CONFLICT        950
FT                   /note="A -> T (in Ref. 4; CAB10973)"
FT                   /evidence="ECO:0000305"
FT   CONFLICT        1137
FT                   /note="V -> I (in Ref. 4; CAB10973)"
FT                   /evidence="ECO:0000305"
FT   CONFLICT        1270..1271
FT                   /note="ST -> RL (in Ref. 1; AAA91230)"
FT                   /evidence="ECO:0000305"
FT   CONFLICT        1276
FT                   /note="P -> S (in Ref. 4; CAB10973)"
FT                   /evidence="ECO:0000305"
FT   CONFLICT        1708
FT                   /note="T -> S (in Ref. 4; CAB10973)"
FT                   /evidence="ECO:0000305"
SQ   SEQUENCE   1741 AA;  185156 MW;  BD770C27B824DE93 CRC64;
     MSRWGKNIVV PLDSLCKEKE NTNRPTVARS VGTVGKWGKM GFTSTRTYTL PAIHPMAAAA
     AAAAAAASPS QSPASTQDHD PNDLSVSVPE PPKPKKFFKS RNTAPPEVIA QIIQQLPHCG
     AGASPMRDHF SSAGAGAGGL TPTSGAQEAG GVKLKPGKGA SSAERKRKSP KKKAATTSAS
     TPSTPGAFYG ASDRDGDGLS DPASEQPEQP SSASGKQKQK KPKEEKKLKP EAPPSRVLGR
     ARKAVNYREV DEDERYPTPT KDLIIPKAGR QPAEVAATAT LAAASSEAFI SSTFGSPGSE
     PSLPPPTSAP SASASTSSQL PSASGSASNP PSASRTPEHP PIVLRISKGT SRLVSTDSEE
     PPSSSPAHQN QLNQLSVTEE EPAERSGDET VPASTPKITV KPLRPPTAAD SVDGSSAAVG
     GASAGDSFEE RKSQSLEPNE DEEEEEEEED EEEEPPEINY CTVKISPDKP PKERLKLIIK
     TDVIRNAIAK AAAAAESRSE KKSRSKKHKH KQLLAAGSGA APASGATPAE INSEFKTPSP
     HLALSEANSQ QAQHTPSHLH QLHQLHPQRG SAVISPTTRS DHDFDSQSSV LGSISSKGNS
     TPQLLAQAVQ EDSCVIRSRG SSVITSDLET SQHSSLVAPP SDIESRLESM MMTIDGAGTG
     AASAVPETPL QEDILAVLRG EVPRLNGNTD PEPTEEEDQQ QQPKRATRGR GRKANNNVDV
     TPPATETRTR GRAKGADATT AAISPPTGKR NTRGTRGSRK AEQEVDMEVD ETAMTTVPAN
     EEQLEQATLP PRRGRNAAAR ANNNNLASVN NNINKIAANL SAKAEASRLA EGGVAGGAAR
     SYGRKRKNQQ VTQVLQQEPV PEEQETPDAE EEQPTPAKIP HTDHREHSPD HDPDPDPDEL
     SNNSNNSSLQ HDGSSSSPPP RDFKFKDKFK RTLTLDTQGA ANAGAGGAAA AAPPESSGEQ
     RGAVKLVISK KKGSIFKSRA LVPSDQAEQA TVAKRHLYKH SWDAALEANG GGTNSDASNA
     SASGVGVAGA KDHLHHLAAG KSDGDFGDSP SSNNNGSSSA CSSASTLRGD SPALGKISRL
     AGKQGVPATS TSSDAFDLDL EPIAGELDLE RSAAGASAGG TGATTGGGGA TGGGGPVRVD
     RKTKDYYPVV RNVKTAHQIQ EIGEYQEMDD DVEYILDALQ PHNPPATRCL SALQLAAKCM
     MPAFRMHVRA HGVVTKFFKA LSDANKDLSL GLCTSAIMYI LSQEGLNMDL DRDSLELMIN
     LLEADGVGGS TETGHPDRAG YDRNKQKVRE LCEEIKAQGK GTHLNVDSLT VGTLAMETLL
     SLTSKRAGEW FKEDLRKLGG LEHIIKTISD FCRPVIACDT EIDWQPTLLD NMQTVARCLR
     VLENVTQHNE TNQRYMLTSG QGKAVETLCQ LYRLCSRQIM LHPSDGGGSN KEHPGVAMRE
     LLVPVLKVLI NLTHTFNEAQ PSLGAELLGQ RGDVVETSFR LLLLSANYIP DQCVFELSIL
     VLTLLINLCM HTVPNRAALM QAAAPAEYVA DNPPAQGSVS ALQALLEYFY KCEELARLVE
     KNTDAFLESN EKGKKKQEEV EETVNNLVQR AGHHMEHTLK GSYAAILVGN LIADNELYES
     VVRRQLRGNS FKEIIGVLEK YHTFMNLTSS LEAAFVAHMK STKRIIDNFK KRDYIYEHSD
     EHDNPLPLNL ETTAQVLAVG ADASHAATSS TTVGSGSAPS STSATGTTRA PRVYKTYSSH
     R
 
 
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