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P35125

- UBP6_HUMAN

UniProt

P35125 - UBP6_HUMAN

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Protein

Ubiquitin carboxyl-terminal hydrolase 6

Gene

USP6

Organism
Homo sapiens (Human)
Status
Reviewed - Annotation score: 5 out of 5- Experimental evidence at protein leveli

Functioni

Deubiquitinase with an ATP-independent isopeptidase activity, cleaving at the C-terminus of the ubiquitin moiety. Catalyzes its own deubiquitination. In vitro, isoform 2, but not isoform 3, shows deubiquitinating activity. Promotes plasma membrane localization of ARF6 and selectively regulates ARF6-dependent endocytic protein trafficking. Is able to initiate tumorigenesis by inducing the production of matrix metalloproteinases following NF-kappa-B activation.3 Publications

Catalytic activityi

Thiol-dependent hydrolysis of ester, thioester, amide, peptide and isopeptide bonds formed by the C-terminal Gly of ubiquitin (a 76-residue protein attached to proteins as an intracellular targeting signal).

Sites

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Active sitei541 – 5411Nucleophile
Active sitei1328 – 13281Proton acceptorPROSITE-ProRule annotation

GO - Molecular functioni

  1. calmodulin binding Source: UniProtKB
  2. cysteine-type endopeptidase activity Source: UniProtKB
  3. nucleic acid binding Source: ProtInc
  4. Rab GTPase activator activity Source: InterPro
  5. ubiquitin-specific protease activity Source: UniProtKB

GO - Biological processi

  1. cellular protein modification process Source: UniProtKB
  2. protein deubiquitination Source: UniProtKB
  3. regulation of vesicle-mediated transport Source: UniProtKB
  4. ubiquitin-dependent protein catabolic process Source: InterPro
Complete GO annotation...

Keywords - Molecular functioni

Hydrolase, Protease, Thiol protease

Keywords - Biological processi

Ubl conjugation pathway

Keywords - Ligandi

Calmodulin-binding

Protein family/group databases

MEROPSiC19.009.

Names & Taxonomyi

Protein namesi
Recommended name:
Ubiquitin carboxyl-terminal hydrolase 6 (EC:3.4.19.12)
Alternative name(s):
Deubiquitinating enzyme 6
Proto-oncogene TRE-2
Ubiquitin thioesterase 6
Ubiquitin-specific-processing protease 6
Gene namesi
Name:USP6
Synonyms:HRP1, TRE2
OrganismiHomo sapiens (Human)
Taxonomic identifieri9606 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
ProteomesiUP000005640: Chromosome 17

Organism-specific databases

HGNCiHGNC:12629. USP6.

Subcellular locationi

Cell membrane. Cytoplasm. Endosome
Note: Localizes to the plasma membrane and to filamentous structures within the cell corresponding to ARF6 regulated tubular endosomes. Activation of RAC1 and CDC42 can direct the relocalization of USP6 to the plasma membrane in a manner that depends on the integrity of the actin cytoskeleton.

GO - Cellular componenti

  1. cytoplasm Source: UniProtKB
  2. lysosome Source: ProtInc
  3. plasma membrane Source: UniProtKB
  4. recycling endosome Source: UniProtKB
Complete GO annotation...

Keywords - Cellular componenti

Cell membrane, Cytoplasm, Endosome, Membrane

Pathology & Biotechi

Involvement in diseasei

A chromosomal aberration involving USP6 is a common genetic feature of aneurysmal bone cyst, a benign osseous neoplasm. Translocation t(16;17)(q22;p13) with CDH11. The translocation generates a fusion gene in which the strong CDH11 promoter is fused to the entire USP6 coding sequence, resulting in USP6 transcriptional up-regulation.

Mutagenesis

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Mutagenesisi150 – 1501T → R: Does not restore GAP activity in yeast complementation assay. 1 Publication
Mutagenesisi187 – 1871R → Q: Does not restore GAP activity in yeast complementation assay. 1 Publication
Mutagenesisi541 – 5411C → S: Loss of enzyme activity. 1 Publication

Keywords - Diseasei

Proto-oncogene

Organism-specific databases

PharmGKBiPA37254.

PTM / Processingi

Molecule processing

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Chaini1 – 14061406Ubiquitin carboxyl-terminal hydrolase 6PRO_0000080625Add
BLAST

Post-translational modificationi

Monubiquitinated; ubiquitination is calmodulin and calcium dependent.1 Publication

Keywords - PTMi

Ubl conjugation

Proteomic databases

MaxQBiP35125.
PaxDbiP35125.
PRIDEiP35125.

PTM databases

PhosphoSiteiP35125.

Expressioni

Tissue specificityi

Testis specific. Expressed in various cancer cell lines.2 Publications

Gene expression databases

BgeeiP35125.
CleanExiHS_USP6.
ExpressionAtlasiP35125. baseline and differential.
GenevestigatoriP35125.

Organism-specific databases

HPAiHPA011742.

Interactioni

Subunit structurei

Interacts with RAC1 and CDC42. Interacts (via Rab-GAP TBC domain) with ARF6. Interacts with calmodulin (CALM1, CALM2 and/or CALM3); the interaction is calcium-dependent.3 Publications

Binary interactionsi

WithEntry#Exp.IntActNotes
ANKRD44Q8N8A24EBI-954590,EBI-1245329
MYL2P109165EBI-954590,EBI-725770

Protein-protein interaction databases

BioGridi114552. 12 interactions.
IntActiP35125. 5 interactions.
MINTiMINT-2865848.
STRINGi9606.ENSP00000250066.

Structurei

3D structure databases

ProteinModelPortaliP35125.
SMRiP35125. Positions 79-313, 528-713, 1031-1110, 1311-1367.
ModBaseiSearch...
MobiDBiSearch...

Family & Domainsi

Domains and Repeats

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Domaini100 – 292193Rab-GAP TBCPROSITE-ProRule annotationAdd
BLAST
Domaini532 – 1369838USPAdd
BLAST

Domaini

The Rab-GAP TBC domain lacks GTPase activator activity but is necessary for interaction with ARF6.

Sequence similaritiesi

Belongs to the peptidase C19 family.Curated
Contains 1 Rab-GAP TBC domain.PROSITE-ProRule annotation
Contains 1 USP domain.Curated

Phylogenomic databases

eggNOGiCOG5210.
GeneTreeiENSGT00670000097750.
HOGENOMiHOG000154762.
InParanoidiP35125.
KOiK11837.
OMAiHDSNIKN.
OrthoDBiEOG7PP55V.
PhylomeDBiP35125.
TreeFamiTF324190.

Family and domain databases

InterProiIPR018200. Pept_C19ubi-hydrolase_C_CS.
IPR001394. Peptidase_C19_UCH.
IPR000195. Rab-GTPase-TBC_dom.
IPR028889. UCH/PAN2.
[Graphical view]
PfamiPF00566. RabGAP-TBC. 1 hit.
PF00443. UCH. 1 hit.
[Graphical view]
SMARTiSM00164. TBC. 1 hit.
[Graphical view]
SUPFAMiSSF47923. SSF47923. 2 hits.
PROSITEiPS50086. TBC_RABGAP. 1 hit.
PS00972. USP_1. 1 hit.
PS00973. USP_2. 1 hit.
PS50235. USP_3. 1 hit.
[Graphical view]

Sequences (3)i

Sequence statusi: Complete.

This entry describes 3 isoformsi produced by alternative splicing. Align

Isoform 1 (identifier: P35125-1) [UniParc]FASTAAdd to Basket

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.

« Hide

        10         20         30         40         50
MDMVENADSL QAQERKDILM KYDKGHRAGL PEDKGPEPVG INSSIDRFGI
60 70 80 90 100
LHETELPPVT AREAKKIRRE MTRTSKWMEM LGEWETYKHS SKLIDRVYKG
110 120 130 140 150
IPMNIRGPVW SVLLNIQEIK LKNPGRYQIM KERGKRSSEH IHHIDLDVRT
160 170 180 190 200
TLRNHVFFRD RYGAKQRELF YILLAYSEYN PEVGYCRDLS HITALFLLYL
210 220 230 240 250
PEEDAFWALV QLLASERHSL PGFHSPNGGT VQGLQDQQEH VVPKSQPKTM
260 270 280 290 300
WHQDKEGLCG QCASLGCLLR NLIDGISLGL TLRLWDVYLV EGEQVLMPIT
310 320 330 340 350
SIALKVQQKR LMKTSRCGLW ARLRNQFFDT WAMNDDTVLK HLRASTKKLT
360 370 380 390 400
RKQGDLPPPA KREQGSLAPR PVPASRGGKT LCKGYRQAPP GPPAQFQRPI
410 420 430 440 450
CSASPPWASR FSTPCPGGAV REDTYPVGTQ GVPSLALAQG GPQGSWRFLE
460 470 480 490 500
WKSMPRLPTD LDIGGPWFPH YDFEWSCWVR AISQEDQLAT CWQAEHCGEV
510 520 530 540 550
HNKDMSWPEE MSFTANSSKI DRQKVPTEKG ATGLSNLGNT CFMNSSIQCV
560 570 580 590 600
SNTQPLTQYF ISGRHLYELN RTNPIGMKGH MAKCYGDLVQ ELWSGTQKSV
610 620 630 640 650
APLKLRRTIA KYAPKFDGFQ QQDSQELLAF LLDGLHEDLN RVHEKPYVEL
660 670 680 690 700
KDSDGRPDWE VAAEAWDNHL RRNRSIIVDL FHGQLRSQVK CKTCGHISVR
710 720 730 740 750
FDPFNFLSLP LPMDSYMDLE ITVIKLDGTT PVRYGLRLNM DEKYTGLKKQ
760 770 780 790 800
LRDLCGLNSE QILLAEVHDS NIKNFPQDNQ KVQLSVSGFL CAFEIPVPSS
810 820 830 840 850
PISASSPTQI DFSSSPSTNG MFTLTTNGDL PKPIFIPNGM PNTVVPCGTE
860 870 880 890 900
KNFTNGMVNG HMPSLPDSPF TGYIIAVHRK MMRTELYFLS PQENRPSLFG
910 920 930 940 950
MPLIVPCTVH TRKKDLYDAV WIQVSWLARP LPPQEASIHA QDRDNCMGYQ
960 970 980 990 1000
YPFTLRVVQK DGNSCAWCPQ YRFCRGCKID CGEDRAFIGN AYIAVDWHPT
1010 1020 1030 1040 1050
ALHLRYQTSQ ERVVDKHESV EQSRRAQAEP INLDSCLRAF TSEEELGESE
1060 1070 1080 1090 1100
MYYCSKCKTH CLATKKLDLW RLPPFLIIHL KRFQFVNDQW IKSQKIVRFL
1110 1120 1130 1140 1150
RESFDPSAFL VPRDPALCQH KPLTPQGDEL SKPRILAREV KKVDAQSSAG
1160 1170 1180 1190 1200
KEDMLLSKSP SSLSANISSS PKGSPSSSRK SGTSCPSSKN SSPNSSPRTL
1210 1220 1230 1240 1250
GRSKGRLRLP QIGSKNKPSS SKKNLDASKE NGAGQICELA DALSRGHMRG
1260 1270 1280 1290 1300
GSQPELVTPQ DHEVALANGF LYEHEACGNG CGDGYSNGQL GNHSEEDSTD
1310 1320 1330 1340 1350
DQREDTHIKP IYNLYAISCH SGILSGGHYI TYAKNPNCKW YCYNDSSCEE
1360 1370 1380 1390 1400
LHPDEIDTDS AYILFYEQQG IDYAQFLPKI DGKKMADTSS TDEDSESDYE

KYSMLQ
Length:1,406
Mass (Da):158,658
Last modified:July 19, 2004 - v2
Checksum:iD3A6822CEB441DB3
GO
Isoform 2 (identifier: P35125-2) [UniParc]FASTAAdd to Basket

Also known as: 213(ORF2)

The sequence of this isoform differs from the canonical sequence as follows:
     1-317: Missing.
     318-359: GLWARLRNQF...TRKQGDLPPP → MPQRLPHARQ...HRDPKDSRDA

Show »
Length:1,089
Mass (Da):121,943
Checksum:iA660606208ABD9EB
GO
Isoform 3 (identifier: P35125-3) [UniParc]FASTAAdd to Basket

Also known as: 210(ORF1), oncTre210p

The sequence of this isoform differs from the canonical sequence as follows:
     774-786: NFPQDNQKVQLSV → ISPLHHLQMECSP
     787-1406: Missing.

Note: Was shown to be tumorigenic in transfected mice and seems not to act as GTPase activating protein.

Show »
Length:786
Mass (Da):89,577
Checksum:i6D5D93D2CE65D879
GO

Experimental Info

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Sequence conflicti963 – 9631N → I in CAA45111. (PubMed:1565468)Curated

Natural variant

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Natural varianti475 – 4751W → R.1 Publication
Corresponds to variant rs8073787 [ dbSNP | Ensembl ].
VAR_051522
Natural varianti525 – 5251V → I.
Corresponds to variant rs2304449 [ dbSNP | Ensembl ].
VAR_059749
Natural varianti912 – 9121R → Q.1 Publication
Corresponds to variant rs9899177 [ dbSNP | Ensembl ].
VAR_051523
Natural varianti1330 – 13301I → V.
Corresponds to variant rs1053611 [ dbSNP | Ensembl ].
VAR_051524

Alternative sequence

Feature keyPosition(s)LengthDescriptionGraphical viewFeature identifierActions
Alternative sequencei1 – 317317Missing in isoform 2. 1 PublicationVSP_010878Add
BLAST
Alternative sequencei318 – 35942GLWAR…DLPPP → MPQRLPHARQHTPLPLGSAD YRRVVSVRPQGPHRDPKDSR DA in isoform 2. 1 PublicationVSP_010879Add
BLAST
Alternative sequencei774 – 78613NFPQD…VQLSV → ISPLHHLQMECSP in isoform 3. 1 PublicationVSP_010880Add
BLAST
Alternative sequencei787 – 1406620Missing in isoform 3. 1 PublicationVSP_010881Add
BLAST

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
X63546 mRNA. Translation: CAA45108.1.
X63547 mRNA. Translation: CAA45111.1.
AY143550 mRNA. Translation: AAN38838.1.
AY163314 Genomic DNA. Translation: AAO21348.1.
CCDSiCCDS11069.2. [P35125-1]
PIRiS57867.
S57868. S22158.
S57874. S22155.
RefSeqiNP_004496.2. NM_004505.2.
XP_005256902.1. XM_005256845.2.
UniGeneiHs.448851.

Genome annotation databases

EnsembliENST00000250066; ENSP00000250066; ENSG00000129204. [P35125-1]
ENST00000572949; ENSP00000461581; ENSG00000129204. [P35125-3]
ENST00000574788; ENSP00000460380; ENSG00000129204. [P35125-1]
GeneIDi9098.
KEGGihsa:9098.
UCSCiuc002gau.1. human. [P35125-1]
uc010ckz.1. human. [P35125-2]

Polymorphism databases

DMDMi50403738.

Keywords - Coding sequence diversityi

Alternative splicing, Chromosomal rearrangement, Polymorphism

Cross-referencesi

Web resourcesi

Atlas of Genetics and Cytogenetics in Oncology and Haematology

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
X63546 mRNA. Translation: CAA45108.1 .
X63547 mRNA. Translation: CAA45111.1 .
AY143550 mRNA. Translation: AAN38838.1 .
AY163314 Genomic DNA. Translation: AAO21348.1 .
CCDSi CCDS11069.2. [P35125-1 ]
PIRi S57867.
S57868. S22158.
S57874. S22155.
RefSeqi NP_004496.2. NM_004505.2.
XP_005256902.1. XM_005256845.2.
UniGenei Hs.448851.

3D structure databases

ProteinModelPortali P35125.
SMRi P35125. Positions 79-313, 528-713, 1031-1110, 1311-1367.
ModBasei Search...
MobiDBi Search...

Protein-protein interaction databases

BioGridi 114552. 12 interactions.
IntActi P35125. 5 interactions.
MINTi MINT-2865848.
STRINGi 9606.ENSP00000250066.

Protein family/group databases

MEROPSi C19.009.

PTM databases

PhosphoSitei P35125.

Polymorphism databases

DMDMi 50403738.

Proteomic databases

MaxQBi P35125.
PaxDbi P35125.
PRIDEi P35125.

Protocols and materials databases

DNASUi 9098.
Structural Biology Knowledgebase Search...

Genome annotation databases

Ensembli ENST00000250066 ; ENSP00000250066 ; ENSG00000129204 . [P35125-1 ]
ENST00000572949 ; ENSP00000461581 ; ENSG00000129204 . [P35125-3 ]
ENST00000574788 ; ENSP00000460380 ; ENSG00000129204 . [P35125-1 ]
GeneIDi 9098.
KEGGi hsa:9098.
UCSCi uc002gau.1. human. [P35125-1 ]
uc010ckz.1. human. [P35125-2 ]

Organism-specific databases

CTDi 9098.
GeneCardsi GC17P005019.
H-InvDB HIX0202554.
HGNCi HGNC:12629. USP6.
HPAi HPA011742.
MIMi 604334. gene.
neXtProti NX_P35125.
PharmGKBi PA37254.
GenAtlasi Search...

Phylogenomic databases

eggNOGi COG5210.
GeneTreei ENSGT00670000097750.
HOGENOMi HOG000154762.
InParanoidi P35125.
KOi K11837.
OMAi HDSNIKN.
OrthoDBi EOG7PP55V.
PhylomeDBi P35125.
TreeFami TF324190.

Miscellaneous databases

ChiTaRSi USP6. human.
GeneWikii USP6.
GenomeRNAii 9098.
NextBioi 34099.
PROi P35125.
SOURCEi Search...

Gene expression databases

Bgeei P35125.
CleanExi HS_USP6.
ExpressionAtlasi P35125. baseline and differential.
Genevestigatori P35125.

Family and domain databases

InterProi IPR018200. Pept_C19ubi-hydrolase_C_CS.
IPR001394. Peptidase_C19_UCH.
IPR000195. Rab-GTPase-TBC_dom.
IPR028889. UCH/PAN2.
[Graphical view ]
Pfami PF00566. RabGAP-TBC. 1 hit.
PF00443. UCH. 1 hit.
[Graphical view ]
SMARTi SM00164. TBC. 1 hit.
[Graphical view ]
SUPFAMi SSF47923. SSF47923. 2 hits.
PROSITEi PS50086. TBC_RABGAP. 1 hit.
PS00972. USP_1. 1 hit.
PS00973. USP_2. 1 hit.
PS50235. USP_3. 1 hit.
[Graphical view ]
ProtoNeti Search...

Publicationsi

« Hide 'large scale' publications
  1. "A novel transcriptional unit of the tre oncogene widely expressed in human cancer cells."
    Nakamura T., Hillova J., Mariage-Samson R., Onno M., Huebner K., Cannizzaro L.A., Boghosian-Sell L., Croce C.M., Hill M.
    Oncogene 7:733-741(1992) [PubMed] [Europe PMC] [Abstract]
    Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORMS 2 AND 3), TISSUE SPECIFICITY, VARIANTS ARG-475 AND GLN-912.
    Tissue: Ewing sarcoma.
  2. Cited for: NUCLEOTIDE SEQUENCE [GENOMIC DNA / MRNA] (ISOFORM 1), ALTERNATIVE SPLICING, TISSUE SPECIFICITY, DISCUSSION OF TRE2 EVOLUTION.
  3. "The yeast DOA4 gene encodes a deubiquitinating enzyme related to a product of the human tre-2 oncogene."
    Papa F.R., Hochstrasser M.
    Nature 366:313-319(1993) [PubMed] [Europe PMC] [Abstract]
    Cited for: CHARACTERIZATION.
  4. "Expression in a RabGAP yeast mutant of two human homologues, one of which is an oncogene."
    Bizimungu C., De Neve N., Burny A., Bach S., Bontemps F., Portetelle D., Vandenbol M.
    Biochem. Biophys. Res. Commun. 310:498-504(2003) [PubMed] [Europe PMC] [Abstract]
    Cited for: MUTAGENESIS OF THR-150 AND ARG-187.
  5. "The TRE17 oncogene encodes a component of a novel effector pathway for Rho GTPases Cdc42 and Rac1 and stimulates actin remodeling."
    Masuda-Robens J.M., Kutney S.N., Qi H., Chou M.M.
    Mol. Cell. Biol. 23:2151-2161(2003) [PubMed] [Europe PMC] [Abstract]
    Cited for: SUBCELLULAR LOCATION, INTERACTION WITH RAC1 AND CDC42.
  6. Cited for: CHROMOSOMAL TRANSLOCATION WITH CDH11.
  7. "The TBC (Tre-2/Bub2/Cdc16) domain protein TRE17 regulates plasma membrane-endosomal trafficking through activation of Arf6."
    Martinu L., Masuda-Robens J.M., Robertson S.E., Santy L.C., Casanova J.E., Chou M.M.
    Mol. Cell. Biol. 24:9752-9762(2004) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, SUBCELLULAR LOCATION, INTERACTION WITH ARF6.
  8. "Calcium/calmodulin regulates ubiquitination of the ubiquitin-specific protease TRE17/USP6."
    Shen C., Ye Y., Robertson S.E., Lau A.W., Mak D.O., Chou M.M.
    J. Biol. Chem. 280:35967-35973(2005) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION, INTERACTION WITH CALMODULIN, UBIQUITINATION, MUTAGENESIS OF CYS-541.
  9. Cited for: IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS].
    Tissue: Cervix carcinoma.
  10. "TRE17/USP6 oncogene translocated in aneurysmal bone cyst induces matrix metalloproteinase production via activation of NF-kappaB."
    Ye Y., Pringle L.M., Lau A.W., Riquelme D.N., Wang H., Jiang T., Lev D., Welman A., Blobel G.A., Oliveira A.M., Chou M.M.
    Oncogene 29:3619-3629(2010) [PubMed] [Europe PMC] [Abstract]
    Cited for: FUNCTION.

Entry informationi

Entry nameiUBP6_HUMAN
AccessioniPrimary (citable) accession number: P35125
Secondary accession number(s): Q15634, Q86WP6, Q8IWT4
Entry historyi
Integrated into UniProtKB/Swiss-Prot: February 1, 1994
Last sequence update: July 19, 2004
Last modified: October 29, 2014
This is version 123 of the entry and version 2 of the sequence. [Complete history]
Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programChordata Protein Annotation Program
DisclaimerAny medical or genetic information present in this entry is provided for research, educational and informational purposes only. It is not in any way intended to be used as a substitute for professional medical advice, diagnosis, treatment or care.

Miscellaneousi

Miscellaneous

The USP6 gene only exists in the primate lineage.

Keywords - Technical termi

Complete proteome, Reference proteome

Documents

  1. Human chromosome 17
    Human chromosome 17: entries, gene names and cross-references to MIM
  2. Human entries with polymorphisms or disease mutations
    List of human entries with polymorphisms or disease mutations
  3. Human polymorphisms and disease mutations
    Index of human polymorphisms and disease mutations
  4. MIM cross-references
    Online Mendelian Inheritance in Man (MIM) cross-references in UniProtKB/Swiss-Prot
  5. Peptidase families
    Classification of peptidase families and list of entries
  6. SIMILARITY comments
    Index of protein domains and families

External Data

Dasty 3