Putative ATP-dependent RNA helicase TDRD9

Preferred order of sections

Names and origin

Protein names

Recommended name:
Putative ATP-dependent RNA helicase TDRD9
EC=3.6.4.13

Alternative name(s):
Tudor domain-containing protein 9

Gene names

Name:

Tdrd9

Organism

Mus musculus (Mouse) [Reference proteome]

Taxonomic identifier

10090 [NCBI]

Taxonomic lineage

Eukaryota › Metazoa › Chordata › Craniata › Vertebrata › Euteleostomi › Mammalia › Eutheria › Euarchontoglires › Glires › Rodentia › Sciurognathi › Muroidea › Muridae › Murinae › Mus › Mus

Protein attributes

Sequence length	1383 AA.
Sequence status	Complete.
Protein existence	Evidence at protein level

General annotation (Comments)

Function	Probable ATP-binding RNA helicase which plays a central role during spermatogenesis by repressing transposable elements and prevent their mobilization, which is essential for the germline integrity. Acts via the piRNA metabolic process, which mediates the repression of transposable elements during meiosis by forming complexes composed of piRNAs and Piwi proteins and govern the methylation and subsequent repression of transposons. Its association with PIWIL4 and the piP-bodies suggests a participation in the secondary piRNAs metabolic process. Ref.4
Catalytic activity	ATP + H₂O = ADP + phosphate.
Subunit structure	Interacts with piRNA-associated proteins PIWIL1 and PIWIL4. Ref.4 Ref.5
Subcellular location	Cytoplasm. Nucleus. Note: Component of the nuage, also named P granule, a germ-cell-specific organelle required to repress transposon during meiosis. Specifically localizes to piP-bodies, a subset of the nuage which contains secondary piRNAs. PIWIL2 is required for its localization to piP-bodies. Ref.4 Ref.6
Tissue specificity	Predominantly expressed in reproductive organs. Detected in mitotic spermatogonia, meiotic spermatocytes (predominantly at the pachytene stage), haploid spermatids in the testis, and in growing oocytes in the ovary (at protein level). Ref.4
Disruption phenotype	Mice are viable but show male sterility with chromosome synapsis failure. In fetal testes, LINE-1 (L1) transposable elements derepression and an aberrant piRNA profile in prospermatogonia, followed by cognate DNA demethylation are observed. Ref.4
Sequence similarities	Belongs to the DEAD box helicase family. DEAH subfamily. Contains 1 helicase ATP-binding domain. Contains 1 helicase C-terminal domain. Contains 1 Tudor domain.

Ontologies

Keywords
Biological process	Differentiation Meiosis RNA-mediated gene silencing Spermatogenesis
Cellular component	Cytoplasm Nucleus
Coding sequence diversity	Alternative splicing
Ligand	ATP-binding Nucleotide-binding
Molecular function	Developmental protein Helicase Hydrolase
Technical term	Complete proteome Reference proteome
Gene Ontology (GO)
Biological_process	DNA methylation involved in gamete generation Inferred from mutant phenotype Ref.4. Source: UniProtKB cell differentiation Inferred from electronic annotation. Source: UniProtKB-KW fertilization Inferred from mutant phenotype Ref.4. Source: UniProtKB gene silencing by RNA Inferred from electronic annotation. Source: UniProtKB-KW male meiosis Inferred from mutant phenotype Ref.4. Source: UniProtKB multicellular organismal development Inferred from electronic annotation. Source: UniProtKB-KW piRNA metabolic process Inferred from mutant phenotype Ref.4. Source: UniProtKB spermatogenesis Inferred from mutant phenotype Ref.4. Source: UniProtKB
Cellular_component	nucleus Inferred from direct assay Ref.4. Source: UniProtKB piP-body Inferred from direct assay Ref.6 Ref.4. Source: UniProtKB
Molecular_function	ATP binding Inferred from electronic annotation. Source: UniProtKB-KW ATP-dependent helicase activity Inferred from electronic annotation. Source: InterPro nucleic acid binding Inferred from electronic annotation. Source: InterPro
Complete GO annotation...

Alternative products

This entry describes 3 isoforms produced by alternative splicing. [Align] [Select]

Isoform 1 (identifier: Q14BI7-1) This isoform has been chosen as the 'canonical' sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

Isoform 2 (identifier: Q14BI7-2) The sequence of this isoform differs from the canonical sequence as follows: 1-401: Missing.

Isoform 3 (identifier: Q14BI7-3) The sequence of this isoform differs from the canonical sequence as follows: 1-772: Missing. 773-813: PKTTVVLKHI...KSIVFDGAKA → MDIGTKCTSQ...SEESCGVPRA
Note: No experimental confirmation available.

Sequence annotation (Features)

Feature key

Position(s)

Length

Description

Graphical view

Feature identifier

Molecule processing

Chain

1 – 1383

1383

Putative ATP-dependent RNA helicase TDRD9

PRO_0000333814

Regions

Domain

144 – 310

167

Helicase ATP-binding

Domain

378 – 545

168

Helicase C-terminal

Domain

945 – 1005

61

Tudor

Nucleotide binding

157 – 164

8

ATP By similarity

Motif

256 – 259

4

DEAH box

Natural variations

Alternative sequence

1 – 772

772

Missing in isoform 3.

VSP_033553

Alternative sequence

1 – 401

401

Missing in isoform 2.

VSP_033554

Alternative sequence

773 – 813

41

PKTTV…DGAKA → MDIGTKCTSQVAAGVTAWHL WRRSGPGRLADSEESCGVPR A in isoform 3.

VSP_033555

Sequences

Sequence		Length	Mass (Da)
Isoform 1 [UniParc]. Last modified March 2, 2010. Version 3. Checksum: 55946F4F639B5777 Show »	FASTA	1,383	155,980
10 20 30 40 50 60 MLRKLTVDQI NDWFTIGKTV TNVELLGLPP AFPAEAPREE VQRSEEVPNE DPTAQAQVPV 70 80 90 100 110 120 KATAPARPAS TSGRSLSQRS SEMEYINKYR QLEEQELDIY GQDQPPSGPG LRSPLAKLSN 130 140 150 160 170 180 VACIPETTYK YPDLPINRCK EEVISLIESN SVVIIHGATG SGKSTQLPQY VLDHYTQRSA 190 200 210 220 230 240 FCNIVVTQPR KIGASSIARW ISKERSWTLG GLVGYQVGLE KIATEDTRLI YMTTGVLLQK 250 260 270 280 290 300 IVSAKSLMEF THIFIDEVHE RTEEMDFLLL VVRKLLRTNS RFVKVVLMSA TINCKQFADY 310 320 330 340 350 360 FAVPVQNKMN PAYVFEVEGK PHAIEEYYLN DLGHIYHSGL PYRLEEPVIT KDVYEVAVSL 370 380 390 400 410 420 IQMFDDLDMK ESGNKTWSGA QFVSERSSVL VFLPGLGEIN YMHELLTNMI HKRLQVYPLH 430 440 450 460 470 480 SSVTLEEQNN VFLSPVPGYR KIILSTNIAE SSVTVPDVKY VIDFCLTRTL VCDEDTNYQS 490 500 510 520 530 540 LRLSWASKTS CDQRKGRAGR VSKGYCYRLI PRDFWDSAIP DHVVPEMLRC PLGSTILKVK 550 560 570 580 590 600 LLDMGEPRAL LATALSPPSL SDIERTILLL KEVGALAVSG QREDENPHDG ELTFLGRVLA 610 620 630 640 650 660 QLPVSQQLGK LVVLGHVFGC LDECLIIAAA LSLKNFFTMP FRQHLDGYRN KVHFSGSSRS 670 680 690 700 710 720 DCLALVEAFR AWQACRQRGE LRRPKDELDW GRLNYIQIKR IREVAELYEE LKNRISQFNM 730 740 750 760 770 780 FVGPHHPVLD QEYPYKQRFI LQVVLAGAFY PNYFTFGQPD EEMAVRELAG KDPKTTVVLK 790 800 810 820 830 840 HIPPYGFLYY KQLQSLFRQC GQVKSIVFDG AKAFVEFSRN PTERFKTLPA VNLAVKMSQL 850 860 870 880 890 900 KVSLELSVHA AEEIEGKVQG GSVSKLRNTR VNVDFQKQTV DPMQVSFNTL DRPRTVADLL 910 920 930 940 950 960 LTIDVTEVVE VGHFWGYRID ERNAELLKQL TAEINRLELV PLPIHPHPDL VCLAPFTDYN 970 980 990 1000 1010 1020 KESYFRAQIL YVSGNSAEVF FVDYGNRSHV DLDLLREIPC QFLELPFQAL EFKICKMRPS 1030 1040 1050 1060 1070 1080 AKSLICGEHW SGGAHGRFAA LVGGCPLLVK VFSIVHSVLH VDVYRYSGAQ DAVNVRDVLI 1090 1100 1110 1120 1130 1140 REGYAELAEE SYESKQSYEV LKGFFAKSVD TMPDGSVSSP LKDDEKHLLR ILLESFASNR 1150 1160 1170 1180 1190 1200 LGAPNCKAVL HGPFNPYELK CHSLTRISKF RCVWIEKESI NSVVISDSPA DLHQRMLVAA 1210 1220 1230 1240 1250 1260 SLSVNETGST MLLRETSLMP HIPGLPALLS MLFAPVMELR VDREGKCYTG VLCGLGWNSA 1270 1280 1290 1300 1310 1320 TEAPILPEHD IELAFDVRLN VEDIVEINIL RAAINKLVCD GPNGSKYLGP ERIAQLQENA 1330 1340 1350 1360 1370 1380 RQKLLGLFCR LKPREKITPQ WHEKPYEWNQ VDPRLIMEQA EPEGSPGKST SLYQLHTPVV LSP « Hide
Isoform 2 [UniParc]. Checksum: 0F9C78208DD4792F Show »	FASTA	982	110,899
10 20 30 40 50 60 MHELLTNMIH KRLQVYPLHS SVTLEEQNNV FLSPVPGYRK IILSTNIAES SVTVPDVKYV 70 80 90 100 110 120 IDFCLTRTLV CDEDTNYQSL RLSWASKTSC DQRKGRAGRV SKGYCYRLIP RDFWDSAIPD 130 140 150 160 170 180 HVVPEMLRCP LGSTILKVKL LDMGEPRALL ATALSPPSLS DIERTILLLK EVGALAVSGQ 190 200 210 220 230 240 REDENPHDGE LTFLGRVLAQ LPVSQQLGKL VVLGHVFGCL DECLIIAAAL SLKNFFTMPF 250 260 270 280 290 300 RQHLDGYRNK VHFSGSSRSD CLALVEAFRA WQACRQRGEL RRPKDELDWG RLNYIQIKRI 310 320 330 340 350 360 REVAELYEEL KNRISQFNMF VGPHHPVLDQ EYPYKQRFIL QVVLAGAFYP NYFTFGQPDE 370 380 390 400 410 420 EMAVRELAGK DPKTTVVLKH IPPYGFLYYK QLQSLFRQCG QVKSIVFDGA KAFVEFSRNP 430 440 450 460 470 480 TERFKTLPAV NLAVKMSQLK VSLELSVHAA EEIEGKVQGG SVSKLRNTRV NVDFQKQTVD 490 500 510 520 530 540 PMQVSFNTLD RPRTVADLLL TIDVTEVVEV GHFWGYRIDE RNAELLKQLT AEINRLELVP 550 560 570 580 590 600 LPIHPHPDLV CLAPFTDYNK ESYFRAQILY VSGNSAEVFF VDYGNRSHVD LDLLREIPCQ 610 620 630 640 650 660 FLELPFQALE FKICKMRPSA KSLICGEHWS GGAHGRFAAL VGGCPLLVKV FSIVHSVLHV 670 680 690 700 710 720 DVYRYSGAQD AVNVRDVLIR EGYAELAEES YESKQSYEVL KGFFAKSVDT MPDGSVSSPL 730 740 750 760 770 780 KDDEKHLLRI LLESFASNRL GAPNCKAVLH GPFNPYELKC HSLTRISKFR CVWIEKESIN 790 800 810 820 830 840 SVVISDSPAD LHQRMLVAAS LSVNETGSTM LLRETSLMPH IPGLPALLSM LFAPVMELRV 850 860 870 880 890 900 DREGKCYTGV LCGLGWNSAT EAPILPEHDI ELAFDVRLNV EDIVEINILR AAINKLVCDG 910 920 930 940 950 960 PNGSKYLGPE RIAQLQENAR QKLLGLFCRL KPREKITPQW HEKPYEWNQV DPRLIMEQAE 970 980 PEGSPGKSTS LYQLHTPVVL SP « Hide
Isoform 3 [UniParc]. Checksum: 43784340033630A0 Show »	FASTA	611	68,306
10 20 30 40 50 60 MDIGTKCTSQ VAAGVTAWHL WRRSGPGRLA DSEESCGVPR AFVEFSRNPT ERFKTLPAVN 70 80 90 100 110 120 LAVKMSQLKV SLELSVHAAE EIEGKVQGGS VSKLRNTRVN VDFQKQTVDP MQVSFNTLDR 130 140 150 160 170 180 PRTVADLLLT IDVTEVVEVG HFWGYRIDER NAELLKQLTA EINRLELVPL PIHPHPDLVC 190 200 210 220 230 240 LAPFTDYNKE SYFRAQILYV SGNSAEVFFV DYGNRSHVDL DLLREIPCQF LELPFQALEF 250 260 270 280 290 300 KICKMRPSAK SLICGEHWSG GAHGRFAALV GGCPLLVKVF SIVHSVLHVD VYRYSGAQDA 310 320 330 340 350 360 VNVRDVLIRE GYAELAEESY ESKQSYEVLK GFFAKSVDTM PDGSVSSPLK DDEKHLLRIL 370 380 390 400 410 420 LESFASNRLG APNCKAVLHG PFNPYELKCH SLTRISKFRC VWIEKESINS VVISDSPADL 430 440 450 460 470 480 HQRMLVAASL SVNETGSTML LRETSLMPHI PGLPALLSML FAPVMELRVD REGKCYTGVL 490 500 510 520 530 540 CGLGWNSATE APILPEHDIE LAFDVRLNVE DIVEINILRA AINKLVCDGP NGSKYLGPER 550 560 570 580 590 600 IAQLQENARQ KLLGLFCRLK PREKITPQWH EKPYEWNQVD PRLIMEQAEP EGSPGKSTSL 610 YQLHTPVVLS P « Hide

References

	« Hide 'large scale' referencesShow 'large scale' references »
[1]	"Regulation of retroelement expression and genome dna methylation through conserved TDRD9/SPN-E function in the germline." Shoji M., Tanaka T., Kitamura K., Hosokawa M., Kato Y., Kondoh G., Okawa K., Sasaki H., Chuma S., Nakatsuji N. Submitted (SEP-2007) to the EMBL/GenBank/DDBJ databases Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1). Tissue: Testis.
[2]	"Lineage-specific biology revealed by a finished genome assembly of the mouse." Church D.M., Goodstadt L., Hillier L.W., Zody M.C., Goldstein S., She X., Bult C.J., Agarwala R., Cherry J.L., DiCuccio M., Hlavina W., Kapustin Y., Meric P., Maglott D., Birtle Z., Marques A.C., Graves T., Zhou S. Ponting C.P. PLoS Biol. 7:E1000112-E1000112(2009) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]. Strain: C57BL/6J.
[3]	"The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC)." The MGC Project Team Genome Res. 14:2121-2127(2004) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORMS 2 AND 3).
[4]	"The TDRD9-MIWI2 complex is essential for piRNA-mediated retrotransposon silencing in the mouse male germline." Shoji M., Tanaka T., Hosokawa M., Reuter M., Stark A., Kato Y., Kondoh G., Okawa K., Chujo T., Suzuki T., Hata K., Martin S.L., Noce T., Kuramochi-Miyagawa S., Nakano T., Sasaki H., Pillai R.S., Nakatsuji N., Chuma S. Dev. Cell 17:775-787(2009) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, SUBCELLULAR LOCATION, DISRUPTION PHENOTYPE, TISSUE SPECIFICITY, INTERACTION WITH PIWIL4.
[5]	"Proteomic analysis of murine Piwi proteins reveals a role for arginine methylation in specifying interaction with Tudor family members." Vagin V.V., Wohlschlegel J., Qu J., Jonsson Z., Huang X., Chuma S., Girard A., Sachidanandam R., Hannon G.J., Aravin A.A. Genes Dev. 23:1749-1762(2009) [PubMed] [Europe PMC] [Abstract] Cited for: INTERACTION WITH PIWIL1 AND PIWIL4.
[6]	"Cytoplasmic compartmentalization of the fetal piRNA pathway in mice." Aravin A.A., van der Heijden G.W., Castaneda J., Vagin V.V., Hannon G.J., Bortvin A. PLoS Genet. 5:E1000764-E1000764(2009) [PubMed] [Europe PMC] [Abstract] Cited for: SUBCELLULAR LOCATION.
+	Additional computationally mapped references.

Cross-references

Sequence databases
EMBL GenBank DDBJ	AB362563 mRNA. Translation: BAG15992.1. AC112520 Genomic DNA. No translation available. AC132623 Genomic DNA. No translation available. BC115831 mRNA. Translation: AAI15832.1. BC116656 mRNA. Translation: AAI16657.1.
IPI	IPI00749982. IPI00849630. IPI00914159.
RefSeq	NP_083332.1. NM_029056.1.
UniGene	Mm.60648.
3D structure databases
ProteinModelPortal	Q14BI7.
SMR	Q14BI7. Positions 126-720, 918-1082.
ModBase	Search...
PTM databases
PhosphoSite	Q14BI7.
Proteomic databases
PaxDb	Q14BI7.
PRIDE	Q14BI7.
Protocols and materials databases
StructuralBiologyKnowledgebase	Search...
Genome annotation databases
Ensembl	ENSMUST00000079009; ENSMUSP00000078022; ENSMUSG00000054003.
GeneID	74691.
KEGG	mmu:74691.
UCSC	uc007pei.1. mouse. uc007pej.1. mouse. uc011yva.1. mouse.
Organism-specific databases
CTD	122402.
MGI	MGI:1921941. Tdrd9.
Phylogenomic databases
eggNOG	COG1643.
GeneTree	ENSGT00700000104393.
HOGENOM	HOG000047965.
HOVERGEN	HBG072266.
InParanoid	B1Q3J8.
OMA	INDWFTI.
OrthoDB	EOG4F4S96.
Gene expression databases
Bgee	Q14BI7.
Genevestigator	Q14BI7.
Family and domain databases
InterPro	IPR011545. DNA/RNA_helicase_DEAD/DEAH_N. IPR007502. Helicase-assoc_dom. IPR014001. Helicase_ATP-bd. IPR001650. Helicase_C. IPR002999. Tudor. [Graphical view]
Pfam	PF00270. DEAD. 1 hit. PF04408. HA2. 1 hit. PF00271. Helicase_C. 1 hit. PF00567. TUDOR. 1 hit. [Graphical view]
SMART	SM00487. DEXDc. 1 hit. SM00847. HA2. 1 hit. SM00490. HELICc. 1 hit. SM00333. TUDOR. 1 hit. [Graphical view]
PROSITE	PS51192. HELICASE_ATP_BIND_1. 1 hit. PS51194. HELICASE_CTER. 1 hit. PS50304. TUDOR. 1 hit. [Graphical view]
ProtoNet	Search...
Other
NextBio	341402.
SOURCE	Search...

Entry information

Entry name

TDRD9_MOUSE

Accession

Primary (citable) accession number: Q14BI7
Secondary accession number(s): B1Q3J8, Q14AW6

Entry history

Integrated into UniProtKB/Swiss-Prot:	May 20, 2008
Last sequence update:	March 2, 2010
Last modified:	May 1, 2013
This is version 59 of the entry and version 3 of the sequence. [Complete history]

Entry status

Reviewed (UniProtKB/Swiss-Prot)

Annotation program

Chordata Protein Annotation Program

Relevant documents

MGD cross-references

Mouse Genome Database (MGD) cross-references in UniProtKB/Swiss-Prot

SIMILARITY comments

Index of protein domains and families

Preferred order of sections

Names and origin

Protein attributes

General annotation (Comments)

Ontologies

Alternative products

Sequence annotation (Features)

Molecule processing

Regions

Natural variations

Sequences

References

Cross-references

Sequence databases

3D structure databases

PTM databases

Proteomic databases

Protocols and materials databases

Genome annotation databases

Organism-specific databases

Phylogenomic databases

Gene expression databases

Family and domain databases

Other

Entry information

Relevant documents