MAEL_DROME
ID MAEL_DROME Reviewed; 459 AA.
AC Q9VNS0; O17317; Q86BG5;
DT 24-MAR-2009, integrated into UniProtKB/Swiss-Prot.
DT 01-MAY-2000, sequence version 1.
DT 03-AUG-2022, entry version 161.
DE RecName: Full=Protein maelstrom;
GN Name=mael; ORFNames=CG11254;
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]
RP NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM A), SUBCELLULAR LOCATION, AND TISSUE
RP SPECIFICITY.
RX PubMed=9409682; DOI=10.1242/dev.124.22.4661;
RA Clegg N.J., Frost D.M., Larkin M.K., Subrahmanyan L., Bryant Z.,
RA Ruohola-Baker H.;
RT "maelstrom is required for an early step in the establishment of Drosophila
RT oocyte polarity: posterior localization of grk mRNA.";
RL Development 124:4661-4671(1997).
RN [2]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC STRAIN=Berkeley;
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]
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]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM A).
RC STRAIN=Berkeley; TISSUE=Embryo;
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 [5]
RP FUNCTION.
RX PubMed=11277405; DOI=10.1007/s004270000114;
RA Clegg N.J., Findley S.D., Mahowald A.P., Ruohola-Baker H.;
RT "Maelstrom is required to position the MTOC in stage 2-6 Drosophila
RT oocytes.";
RL Dev. Genes Evol. 211:44-48(2001).
RN [6]
RP FUNCTION, SUBCELLULAR LOCATION, AND DISRUPTION PHENOTYPE.
RX PubMed=12538514; DOI=10.1242/dev.00310;
RA Findley S.D., Tamanaha M., Clegg N.J., Ruohola-Baker H.;
RT "Maelstrom, a Drosophila spindle-class gene, encodes a protein that
RT colocalizes with Vasa and RDE1/AGO1 homolog, Aubergine, in nuage.";
RL Development 130:859-871(2003).
RN [7]
RP FUNCTION, SUBCELLULAR LOCATION, AND DISRUPTION PHENOTYPE.
RX PubMed=17428915; DOI=10.1073/pnas.0701920104;
RA Lim A.K., Kai T.;
RT "Unique germ-line organelle, nuage, functions to repress selfish genetic
RT elements in Drosophila melanogaster.";
RL Proc. Natl. Acad. Sci. U.S.A. 104:6714-6719(2007).
RN [8]
RP ERRATUM OF PUBMED:17428915.
RA Lim A.K., Kai T.;
RL Proc. Natl. Acad. Sci. U.S.A. 104:20143-20143(2007).
RN [9]
RP FUNCTION, SUBCELLULAR LOCATION, DNA-BINDING, AND DISRUPTION PHENOTYPE.
RX PubMed=19758565; DOI=10.1016/j.devcel.2009.07.017;
RA Pek J.W., Lim A.K., Kai T.;
RT "Drosophila maelstrom ensures proper germline stem cell lineage
RT differentiation by repressing microRNA-7.";
RL Dev. Cell 17:417-424(2009).
RN [10]
RP FUNCTION, SUBCELLULAR LOCATION, TISSUE SPECIFICITY, DISRUPTION PHENOTYPE,
RP AND MUTAGENESIS OF GLU-131 AND HIS-291.
RX PubMed=23159368; DOI=10.1016/j.cell.2012.10.040;
RA Sienski G., Donertas D., Brennecke J.;
RT "Transcriptional silencing of transposons by Piwi and maelstrom and its
RT impact on chromatin state and gene expression.";
RL Cell 151:964-980(2012).
RN [11]
RP SUBCELLULAR LOCATION.
RX PubMed=23913921; DOI=10.1101/gad.221515.113;
RA Ohtani H., Iwasaki Y.W., Shibuya A., Siomi H., Siomi M.C., Saito K.;
RT "DmGTSF1 is necessary for Piwi-piRISC-mediated transcriptional transposon
RT silencing in the Drosophila ovary.";
RL Genes Dev. 27:1656-1661(2013).
CC -!- FUNCTION: Involved both in the piRNA and miRNA metabolic processes. As
CC a component of the meiotic nuage, plays a central role during oogenesis
CC by repressing transposable elements and preventing their mobilization,
CC which is essential for the germline integrity. Repression of
CC transposable elements is mediated via the piRNA metabolic process,
CC which mediates the repression of transposable elements during meiosis
CC by forming complexes composed of piRNAs and Piwi proteins and governs
CC the repression of transposons. As a nuclear component, it is required
CC for proper differentiation in the germline stem cell (GSC) lineage by
CC repressing microRNA-7 (miR-7), thereby acting as an indirect regulator
CC of bag-of-marbles (Bam). Acts by binding to the promoter of miR-7 gene
CC and repressing its expression; miR-7 repression alleviates the Bam
CC repression by miR-7, thereby allowing differentiation in the germline
CC stem cell (GSC) lineage. Indirectly required to position the
CC microtubule organizing center in stage 2-6 oocytes.
CC {ECO:0000269|PubMed:11277405, ECO:0000269|PubMed:12538514,
CC ECO:0000269|PubMed:17428915, ECO:0000269|PubMed:19758565,
CC ECO:0000269|PubMed:23159368}.
CC -!- SUBCELLULAR LOCATION: Cytoplasm. Nucleus {ECO:0000269|PubMed:23913921}.
CC Note=Component of the meiotic nuage, also named P granule, a germ-cell-
CC specific organelle required to repress transposon activity during
CC meiosis. Vas, aub and spn-E are required for nuage localization.
CC Shuttles between the cytoplasm and the nucleus.
CC -!- ALTERNATIVE PRODUCTS:
CC Event=Alternative splicing; Named isoforms=2;
CC Name=A;
CC IsoId=Q9VNS0-1; Sequence=Displayed;
CC Name=C;
CC IsoId=Q9VNS0-2; Sequence=VSP_036675;
CC -!- TISSUE SPECIFICITY: In germaria and egg chambers, it is detected in the
CC germline. In the germarium, it is in all regions, including region I
CC where the germ cells are dividing. In early egg chambers, it is
CC uniformly distributed throughout the nurse cells and oocyte but, by
CC stage 5, it is most concentrated around the outer margins of the cells,
CC closest to the periphery of the egg chamber. Level decreases in stages
CC 5 and 6, but most noticeably in the oocyte, where protein level
CC remains. No detectable protein from stage 8 onward (at protein level).
CC {ECO:0000269|PubMed:23159368, ECO:0000269|PubMed:9409682}.
CC -!- DISRUPTION PHENOTYPE: Female sterility and defects in karyosome
CC formation and oocyte polarity due to transposable element derepression.
CC Ovary shows mislocalization of 2 proteins involved in the microRNA
CC and/or RNAi pathways, Dicer and AGO2. In testis, transit-amplifying
CC cysts fail to differentiate into primary spermatocytes, instead
CC breaking down into ectopic germline stem cells (GSC) and smaller cysts,
CC due to a depletion of Bag-of-marbles (Bam) protein.
CC {ECO:0000269|PubMed:12538514, ECO:0000269|PubMed:17428915,
CC ECO:0000269|PubMed:19758565, ECO:0000269|PubMed:23159368}.
CC -!- SIMILARITY: Belongs to the maelstrom family. {ECO:0000305}.
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DR EMBL; AF025953; AAB97831.1; -; mRNA.
DR EMBL; AE014296; AAF51851.1; -; Genomic_DNA.
DR EMBL; AE014296; AAF51852.1; -; Genomic_DNA.
DR EMBL; AE014296; AAO41286.1; -; Genomic_DNA.
DR EMBL; AY119598; AAM50252.1; -; mRNA.
DR RefSeq; NP_001303390.1; NM_001316461.1. [Q9VNS0-1]
DR RefSeq; NP_524217.1; NM_079493.4. [Q9VNS0-1]
DR RefSeq; NP_730739.1; NM_168958.3. [Q9VNS0-1]
DR RefSeq; NP_788566.1; NM_176388.3. [Q9VNS0-2]
DR PDB; 4YBG; X-ray; 1.60 A; A=84-333.
DR PDBsum; 4YBG; -.
DR AlphaFoldDB; Q9VNS0; -.
DR SMR; Q9VNS0; -.
DR BioGRID; 65723; 20.
DR IntAct; Q9VNS0; 3.
DR STRING; 7227.FBpp0088574; -.
DR PaxDb; Q9VNS0; -.
DR ABCD; Q9VNS0; 5 sequenced antibodies.
DR DNASU; 40489; -.
DR EnsemblMetazoa; FBtr0089631; FBpp0088574; FBgn0016034. [Q9VNS0-2]
DR EnsemblMetazoa; FBtr0089632; FBpp0088575; FBgn0016034. [Q9VNS0-1]
DR EnsemblMetazoa; FBtr0089633; FBpp0088576; FBgn0016034. [Q9VNS0-1]
DR EnsemblMetazoa; FBtr0347062; FBpp0312456; FBgn0016034. [Q9VNS0-1]
DR GeneID; 40489; -.
DR KEGG; dme:Dmel_CG11254; -.
DR UCSC; CG11254-RA; d. melanogaster. [Q9VNS0-1]
DR UCSC; CG11254-RC; d. melanogaster.
DR CTD; 84944; -.
DR FlyBase; FBgn0016034; mael.
DR VEuPathDB; VectorBase:FBgn0016034; -.
DR eggNOG; ENOG502QTQB; Eukaryota.
DR GeneTree; ENSGT00390000003645; -.
DR InParanoid; Q9VNS0; -.
DR OMA; YANIVEY; -.
DR PhylomeDB; Q9VNS0; -.
DR SignaLink; Q9VNS0; -.
DR BioGRID-ORCS; 40489; 0 hits in 1 CRISPR screen.
DR GenomeRNAi; 40489; -.
DR PRO; PR:Q9VNS0; -.
DR Proteomes; UP000000803; Chromosome 3L.
DR Bgee; FBgn0016034; Expressed in egg chamber and 30 other tissues.
DR ExpressionAtlas; Q9VNS0; baseline and differential.
DR Genevisible; Q9VNS0; DM.
DR GO; GO:0005737; C:cytoplasm; IDA:FlyBase.
DR GO; GO:0005634; C:nucleus; IDA:UniProtKB.
DR GO; GO:0043186; C:P granule; IDA:FlyBase.
DR GO; GO:0048471; C:perinuclear region of cytoplasm; IDA:UniProtKB.
DR GO; GO:0043565; F:sequence-specific DNA binding; IDA:FlyBase.
DR GO; GO:0000976; F:transcription cis-regulatory region binding; IDA:UniProtKB.
DR GO; GO:0007010; P:cytoskeleton organization; TAS:FlyBase.
DR GO; GO:0046843; P:dorsal appendage formation; IMP:FlyBase.
DR GO; GO:0009950; P:dorsal/ventral axis specification; IMP:FlyBase.
DR GO; GO:0031047; P:gene silencing by RNA; IMP:UniProtKB.
DR GO; GO:0030718; P:germ-line stem cell population maintenance; IMP:UniProtKB.
DR GO; GO:0008298; P:intracellular mRNA localization; IMP:FlyBase.
DR GO; GO:0007140; P:male meiotic nuclear division; IBA:GO_Central.
DR GO; GO:0031023; P:microtubule organizing center organization; IMP:UniProtKB.
DR GO; GO:0010586; P:miRNA metabolic process; IMP:UniProtKB.
DR GO; GO:0045892; P:negative regulation of transcription, DNA-templated; IDA:UniProtKB.
DR GO; GO:0010529; P:negative regulation of transposition; IMP:FlyBase.
DR GO; GO:0007314; P:oocyte anterior/posterior axis specification; IMP:FlyBase.
DR GO; GO:0048600; P:oocyte fate commitment; IMP:FlyBase.
DR GO; GO:0016325; P:oocyte microtubule cytoskeleton organization; IMP:FlyBase.
DR GO; GO:0051663; P:oocyte nucleus localization involved in oocyte dorsal/ventral axis specification; IMP:FlyBase.
DR GO; GO:0048477; P:oogenesis; IMP:FlyBase.
DR GO; GO:0034587; P:piRNA metabolic process; IMP:UniProtKB.
DR GO; GO:0008104; P:protein localization; IMP:FlyBase.
DR GO; GO:0060964; P:regulation of miRNA-mediated gene silencing; IDA:UniProtKB.
DR GO; GO:0007317; P:regulation of pole plasm oskar mRNA localization; IMP:FlyBase.
DR GO; GO:0007283; P:spermatogenesis; IMP:UniProtKB.
DR Gene3D; 1.10.30.10; -; 1.
DR InterPro; IPR036910; HMG_box_dom_sf.
DR InterPro; IPR024970; Maelstrom.
DR InterPro; IPR039259; Protein_maelstrom.
DR PANTHER; PTHR21358; PTHR21358; 1.
DR Pfam; PF13017; Maelstrom; 1.
DR SUPFAM; SSF47095; SSF47095; 1.
PE 1: Evidence at protein level;
KW 3D-structure; Alternative splicing; Cytoplasm; Developmental protein;
KW Differentiation; DNA-binding; Meiosis; Nucleus; Oogenesis;
KW Reference proteome; Repressor; RNA-mediated gene silencing; Transcription;
KW Transcription regulation.
FT CHAIN 1..459
FT /note="Protein maelstrom"
FT /id="PRO_0000367299"
FT DNA_BIND 2..69
FT /note="HMG box"
FT REGION 43..78
FT /note="Disordered"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT VAR_SEQ 40
FT /note="E -> EPFQ (in isoform C)"
FT /evidence="ECO:0000305"
FT /id="VSP_036675"
FT MUTAGEN 131
FT /note="E->A: Reduces nuclear aCCumulation in ovary and
FT ovarian somatic cells. Affects transposable element
FT silencing. Causes female sterility."
FT /evidence="ECO:0000269|PubMed:23159368"
FT MUTAGEN 291
FT /note="H->A: Reduces nuclear aCCumulation in ovary and
FT ovarian somatic cells. Affects transposable element
FT silencing. Causes female sterility."
FT /evidence="ECO:0000269|PubMed:23159368"
FT CONFLICT 247
FT /note="L -> F (in Ref. 1; AAB97831)"
FT /evidence="ECO:0000305"
FT HELIX 85..101
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 105..107
FT /evidence="ECO:0007829|PDB:4YBG"
FT STRAND 110..120
FT /evidence="ECO:0007829|PDB:4YBG"
FT STRAND 122..125
FT /evidence="ECO:0007829|PDB:4YBG"
FT STRAND 127..138
FT /evidence="ECO:0007829|PDB:4YBG"
FT TURN 139..141
FT /evidence="ECO:0007829|PDB:4YBG"
FT STRAND 142..151
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 164..170
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 186..201
FT /evidence="ECO:0007829|PDB:4YBG"
FT STRAND 203..205
FT /evidence="ECO:0007829|PDB:4YBG"
FT STRAND 207..211
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 213..215
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 216..226
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 231..234
FT /evidence="ECO:0007829|PDB:4YBG"
FT STRAND 239..243
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 244..258
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 268..277
FT /evidence="ECO:0007829|PDB:4YBG"
FT TURN 279..282
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 289..294
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 297..299
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 301..316
FT /evidence="ECO:0007829|PDB:4YBG"
FT HELIX 317..319
FT /evidence="ECO:0007829|PDB:4YBG"
FT TURN 327..329
FT /evidence="ECO:0007829|PDB:4YBG"
SQ SEQUENCE 459 AA; 51601 MW; 002F7F1298CFEF4E CRC64;
MAPKKHSGFM MFVNEWRNRN AEGRRMTLAQ AVSHCGTIWE KMNTQQRGPY NSGGKDANVA
QRAKRESSNG HGQVDKAQRE ATESLMDMKR TIERLVLNAK MSHDLENAKF VFVAFNYFTK
ALTTDVYVPA EFAACEYSLK EGIRSIYSTM IDPGQIIFGQ GSDALLHSST THDLPLPPNA
LGEKNMTKLY RNIVDYLSKC QGKGKTLVVF TPAENITMVK SCFRYLECDD DFRDGGEKIQ
VFDIQYLLFI LKKEVMNVAD LNDEKINKFA TDAFFKKDFF EFTAGIACQY HEDNDRTKYC
TQSMVTRWAY TFTDFMCGDL AITVQPGKHI PAQTKPNYLI ISSYASSLDH ESSFDSFYSL
PGSGVKKESQ PEACSLSSSR LSVASSSYKP IDHTSFAANL NEVSEFPSLG MRNSSKHHGI
AASAQREWNA RNLPTHSRLI RKVSDNDFSV NGADGKLKK
