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Protein

Drebrin-like protein

Gene

DBNL

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

Functioni

Adapter protein that binds F-actin and DNM1, and thereby plays a role in receptor-mediated endocytosis. Plays a role in the reorganization of the actin cytoskeleton, formation of cell projections, such as neurites, in neuron morphogenesis and synapse formation via its interaction with WASL and COBL. Does not bind G-actin and promote actin polymerization by itself. Required for the formation of organized podosome rosettes (By similarity). May act as a common effector of antigen receptor-signaling pathways in leukocytes. Acts as a key component of the immunological synapse that regulates T-cell activation by bridging TCRs and the actin cytoskeleton to gene activation and endocytic processes.By similarity1 Publication

GO - Molecular functioni

  • actin binding Source: UniProtKB
  • actin filament binding Source: UniProtKB
  • cadherin binding involved in cell-cell adhesion Source: BHF-UCL
  • enzyme activator activity Source: ProtInc

GO - Biological processi

Complete GO annotation...

Keywords - Biological processi

Adaptive immunity, Endocytosis, Immunity, Transport

Keywords - Ligandi

Actin-binding

Enzyme and pathway databases

ReactomeiR-HSA-264870. Caspase-mediated cleavage of cytoskeletal proteins.
R-HSA-6794361. Interactions of neurexins and neuroligins at synapses.
R-HSA-6798695. Neutrophil degranulation.
SignaLinkiQ9UJU6.
SIGNORiQ9UJU6.

Names & Taxonomyi

Protein namesi
Recommended name:
Drebrin-like protein
Alternative name(s):
Cervical SH3P7
Cervical mucin-associated protein
Drebrin-F
HPK1-interacting protein of 55 kDa
Short name:
HIP-55
SH3 domain-containing protein 7
Gene namesi
Name:DBNL
Synonyms:CMAP, SH3P7
ORF Names:PP5423
OrganismiHomo sapiens (Human)
Taxonomic identifieri9606 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
Proteomesi
  • UP000005640 Componenti: Chromosome 7

Organism-specific databases

HGNCiHGNC:2696. DBNL.

Subcellular locationi

  • Cytoplasmcytoskeleton By similarity
  • Cell projectionlamellipodium By similarity
  • Cell projectionruffle By similarity
  • Cytoplasmcell cortex By similarity
  • Cytoplasmcytosol By similarity
  • Cell junctionsynapse By similarity
  • Cell projection By similarity
  • Cell membrane 1 Publication; Peripheral membrane protein; Cytoplasmic side By similarity
  • Cytoplasmic vesicleclathrin-coated vesicle membrane By similarity; Peripheral membrane protein; Cytoplasmic side
  • Golgi apparatus membrane By similarity; Peripheral membrane protein; Cytoplasmic side
  • Cell projectionpodosome By similarity
  • Early endosome 1 Publication

  • Note: Detected in neuron cell body and cell projections, such as neurites. Colocalizes with cytosolic dynamin in hippocampus neurons (By similarity). Cortical actin cytoskeleton. Associates with lamellipodial actin and membrane ruffles. Colocalizes with actin and cortactin at podosome dots and podosome rosettes (By similarity).By similarity

GO - Cellular componenti

Complete GO annotation...

Keywords - Cellular componenti

Cell junction, Cell membrane, Cell projection, Cytoplasm, Cytoplasmic vesicle, Cytoskeleton, Endosome, Golgi apparatus, Membrane, Synapse

Pathology & Biotechi

Mutagenesis

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Mutagenesisi361D → A: Abolishes cleavage by caspase-3. 1 Publication1

Organism-specific databases

DisGeNETi28988.
OpenTargetsiENSG00000136279.
PharmGKBiPA27164.

Polymorphism and mutation databases

BioMutaiDBNL.
DMDMi51316115.

PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
ChainiPRO_00000797931 – 430Drebrin-like proteinAdd BLAST430

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Modified residuei26PhosphothreonineBy similarity1
Modified residuei160PhosphoserineCombined sources1
Modified residuei176N6-acetyllysineCombined sources1
Modified residuei232PhosphoserineCombined sources1
Modified residuei269PhosphoserineCombined sources1
Modified residuei272PhosphoserineCombined sources1
Modified residuei275PhosphoserineCombined sources1
Modified residuei283PhosphoserineCombined sources1
Modified residuei288N6-acetyllysineCombined sources1
Modified residuei291PhosphothreonineCombined sources1
Modified residuei334PhosphotyrosineBy similarity1
Modified residuei344PhosphotyrosineBy similarity1
Isoform 4 (identifier: Q9UJU6-4)
Modified residuei137PhosphoserineCombined sources1
Isoform 6 (identifier: Q9UJU6-6)
Modified residuei183PhosphoserineCombined sources1
Isoform 2 (identifier: Q9UJU6-2)
Modified residuei232PhosphoserineCombined sources1
Isoform 3 (identifier: Q9UJU6-3)
Modified residuei232PhosphoserineCombined sources1

Post-translational modificationi

Degraded by caspases during apoptosis.

Sites

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Sitei361 – 362Cleavage; by caspase-32

Keywords - PTMi

Acetylation, Phosphoprotein

Proteomic databases

EPDiQ9UJU6.
PaxDbiQ9UJU6.
PeptideAtlasiQ9UJU6.
PRIDEiQ9UJU6.
TopDownProteomicsiQ9UJU6-2. [Q9UJU6-2]

2D gel databases

OGPiQ9UJU6.

PTM databases

iPTMnetiQ9UJU6.
PhosphoSitePlusiQ9UJU6.

Miscellaneous databases

PMAP-CutDBQ9UJU6.

Expressioni

Gene expression databases

BgeeiENSG00000136279.
CleanExiHS_DBNL.
ExpressionAtlasiQ9UJU6. baseline and differential.
GenevisibleiQ9UJU6. HS.

Organism-specific databases

HPAiHPA020265.
HPA027735.

Interactioni

Subunit structurei

Interacts with SHANK2, SHANK3 and SYN1. Interacts with FGD1 and DNM1. Interacts with ANKRD54. Interacts with COBL. Interacts with WASL and WIPF1 (By similarity). Interacts with MAP4K1 and PRAM1.By similarity2 Publications

Binary interactionsi

WithEntry#Exp.IntActNotes
ACDQ96AP02EBI-751783,EBI-717666
SH2D4AQ9H7886EBI-751783,EBI-747035
SH2D4AQ9H788-23EBI-751783,EBI-10308083
SH3BP2P783147EBI-751783,EBI-727062

GO - Molecular functioni

Protein-protein interaction databases

BioGridi118809. 50 interactors.
IntActiQ9UJU6. 8 interactors.
MINTiMINT-1474845.
STRINGi9606.ENSP00000417653.

Structurei

Secondary structure

1430
Legend: HelixTurnBeta strandPDB Structure known for this area
Show more details
Feature keyPosition(s)DescriptionActionsGraphical viewLength
Helixi9 – 20Combined sources12
Beta strandi22 – 25Combined sources4
Beta strandi27 – 38Combined sources12
Beta strandi40 – 48Combined sources9
Helixi50 – 56Combined sources7
Beta strandi61 – 70Combined sources10
Beta strandi72 – 74Combined sources3
Beta strandi76 – 85Combined sources10
Helixi91 – 107Combined sources17
Turni108 – 110Combined sources3
Beta strandi111 – 115Combined sources5
Helixi120 – 123Combined sources4
Helixi125 – 133Combined sources9

3D structure databases

Select the link destinations:
PDBei
RCSB PDBi
PDBji
Links Updated
PDB entryMethodResolution (Å)ChainPositionsPDBsum
1X67NMR-A1-133[»]
ProteinModelPortaliQ9UJU6.
SMRiQ9UJU6.
ModBaseiSearch...
MobiDBiSearch...

Miscellaneous databases

EvolutionaryTraceiQ9UJU6.

Family & Domainsi

Domains and Repeats

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Domaini4 – 133ADF-HPROSITE-ProRule annotationAdd BLAST130
Domaini371 – 430SH3PROSITE-ProRule annotationAdd BLAST60

Coiled coil

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Coiled coili176 – 231Sequence analysisAdd BLAST56

Domaini

The SH3 domain mediates interaction with SHANK2, SHANK3 and PRAM1.

Sequence similaritiesi

Belongs to the ABP1 family.Curated
Contains 1 ADF-H domain.PROSITE-ProRule annotation
Contains 1 SH3 domain.PROSITE-ProRule annotation

Keywords - Domaini

Coiled coil, SH3 domain

Phylogenomic databases

eggNOGiKOG3655. Eukaryota.
ENOG410XRVX. LUCA.
GeneTreeiENSGT00530000062953.
HOGENOMiHOG000008567.
HOVERGENiHBG051316.
InParanoidiQ9UJU6.
KOiK20520.
OMAiFQDTGPQ.
OrthoDBiEOG091G05KP.
PhylomeDBiQ9UJU6.
TreeFamiTF318935.

Family and domain databases

Gene3Di3.40.20.10. 1 hit.
InterProiIPR002108. ADF-H.
IPR029006. ADF-H/Gelsolin-like_dom.
IPR029923. Dbnl.
IPR001452. SH3_domain.
[Graphical view]
PANTHERiPTHR10829:SF12. PTHR10829:SF12. 1 hit.
PfamiPF00241. Cofilin_ADF. 1 hit.
PF14604. SH3_9. 1 hit.
[Graphical view]
SMARTiSM00102. ADF. 1 hit.
SM00326. SH3. 1 hit.
[Graphical view]
SUPFAMiSSF50044. SSF50044. 1 hit.
PROSITEiPS51263. ADF_H. 1 hit.
PS50002. SH3. 1 hit.
[Graphical view]

Sequences (6)i

Sequence statusi: Complete.

This entry describes 6 isoformsi produced by alternative splicing. AlignAdd to basket

Isoform 1 (identifier: Q9UJU6-1) [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
MAANLSRNGP ALQEAYVRVV TEKSPTDWAL FTYEGNSNDI RVAGTGEGGL
60 70 80 90 100
EEMVEELNSG KVMYAFCRVK DPNSGLPKFV LINWTGEGVN DVRKGACASH
110 120 130 140 150
VSTMASFLKG AHVTINARAE EDVEPECIME KVAKASGANY SFHKESGRFQ
160 170 180 190 200
DVGPQAPVGS VYQKTNAVSE IKRVGKDSFW AKAEKEEENR RLEEKRRAEE
210 220 230 240 250
AQRQLEQERR ERELREAARR EQRYQEQGGE ASPQRTWEQQ QEVVSRNRNE
260 270 280 290 300
QESAVHPREI FKQKERAMST TSISSPQPGK LRSPFLQKQL TQPETHFGRE
310 320 330 340 350
PAAAISRPRA DLPAEEPAPS TPPCLVQAEE EAVYEEPPEQ ETFYEQPPLV
360 370 380 390 400
QQQGAGSEHI DHHIQGQGLS GQGLCARALY DYQAADDTEI SFDPENLITG
410 420 430
IEVIDEGWWR GYGPDGHFGM FPANYVELIE
Length:430
Mass (Da):48,207
Last modified:May 1, 2000 - v1
Checksum:i7E8C42ED047257AE
GO
Isoform 2 (identifier: Q9UJU6-2) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     234-234: Q → QS

Note: No experimental confirmation available.Combined sources
Show »
Length:431
Mass (Da):48,294
Checksum:iD18F9D316FFD0B4E
GO
Isoform 3 (identifier: Q9UJU6-3) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     234-234: Q → QS
     251-251: Q → QGSTCASLQ

Note: No experimental confirmation available.Combined sources
Show »
Length:439
Mass (Da):49,042
Checksum:i8C044FBD0E82C6D5
GO
Isoform 4 (identifier: Q9UJU6-4) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     1-109: MAANLSRNGP...HVSTMASFLK → MKATAMTSAWLAQG
     234-234: Q → QS

Show »
Length:336
Mass (Da):37,959
Checksum:iCDDA7B4BDFAADB32
GO
Isoform 5 (identifier: Q9UJU6-5) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     1-103: Missing.

Show »
Length:327
Mass (Da):37,101
Checksum:iBE4C189EDD962834
GO
Isoform 6 (identifier: Q9UJU6-6) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     110-158: Missing.
     234-234: Q → QS

Note: No experimental confirmation available.Combined sources
Show »
Length:382
Mass (Da):43,041
Checksum:i9B904F935E7107E7
GO

Experimental Info

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Sequence conflicti8N → K in AAG17262 (PubMed:15498874).Curated1
Sequence conflicti98A → S in AAF81273 (Ref. 3) Curated1
Sequence conflicti98A → S in AAG13120 (Ref. 3) Curated1
Sequence conflicti235R → S in AAF81273 (Ref. 3) Curated1
Sequence conflicti235R → S in AAG13120 (Ref. 3) Curated1
Sequence conflicti430E → D in CAG33448 (Ref. 6) Curated1

Alternative sequence

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Alternative sequenceiVSP_0547791 – 109MAANL…ASFLK → MKATAMTSAWLAQG in isoform 4. CuratedAdd BLAST109
Alternative sequenceiVSP_0547801 – 103Missing in isoform 5. 1 PublicationAdd BLAST103
Alternative sequenceiVSP_057346110 – 158Missing in isoform 6. 1 PublicationAdd BLAST49
Alternative sequenceiVSP_011398234Q → QS in isoform 2, isoform 3, isoform 4 and isoform 6. 2 Publications1
Alternative sequenceiVSP_011399251Q → QGSTCASLQ in isoform 3. 1 Publication1

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AF197060 mRNA. Translation: AAF13701.1.
AF077353 mRNA. Translation: AAF80228.1.
AF250287 mRNA. Translation: AAF81273.1.
AF151364 mRNA. Translation: AAG13120.1.
AF218020 mRNA. Translation: AAG17262.2.
AK027367 mRNA. Translation: BAB55065.1.
AK293279 mRNA. Translation: BAG56807.1.
AK293698 mRNA. Translation: BAG57133.1.
CR457167 mRNA. Translation: CAG33448.1.
AC017116 Genomic DNA. No translation available.
CH236960 Genomic DNA. Translation: EAL23770.1.
CH471128 Genomic DNA. Translation: EAW61132.1.
BC011677 mRNA. Translation: AAH11677.1.
BC031687 mRNA. Translation: AAH31687.1.
CCDSiCCDS34622.1. [Q9UJU6-2]
CCDS34623.1. [Q9UJU6-1]
CCDS47579.1. [Q9UJU6-3]
CCDS64633.1. [Q9UJU6-4]
CCDS64634.1. [Q9UJU6-5]
RefSeqiNP_001014436.1. NM_001014436.2. [Q9UJU6-1]
NP_001116428.1. NM_001122956.1. [Q9UJU6-3]
NP_001271242.1. NM_001284313.1. [Q9UJU6-5]
NP_001271244.1. NM_001284315.1. [Q9UJU6-4]
NP_054782.2. NM_014063.6. [Q9UJU6-2]
UniGeneiHs.436500.

Genome annotation databases

EnsembliENST00000440166; ENSP00000415173; ENSG00000136279. [Q9UJU6-5]
ENST00000448521; ENSP00000411701; ENSG00000136279. [Q9UJU6-1]
ENST00000456905; ENSP00000416421; ENSG00000136279. [Q9UJU6-6]
ENST00000468694; ENSP00000417653; ENSG00000136279. [Q9UJU6-3]
ENST00000490734; ENSP00000417749; ENSG00000136279. [Q9UJU6-4]
ENST00000494774; ENSP00000419992; ENSG00000136279. [Q9UJU6-2]
GeneIDi28988.
KEGGihsa:28988.
UCSCiuc003tjo.5. human. [Q9UJU6-1]

Keywords - Coding sequence diversityi

Alternative splicing

Cross-referencesi

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AF197060 mRNA. Translation: AAF13701.1.
AF077353 mRNA. Translation: AAF80228.1.
AF250287 mRNA. Translation: AAF81273.1.
AF151364 mRNA. Translation: AAG13120.1.
AF218020 mRNA. Translation: AAG17262.2.
AK027367 mRNA. Translation: BAB55065.1.
AK293279 mRNA. Translation: BAG56807.1.
AK293698 mRNA. Translation: BAG57133.1.
CR457167 mRNA. Translation: CAG33448.1.
AC017116 Genomic DNA. No translation available.
CH236960 Genomic DNA. Translation: EAL23770.1.
CH471128 Genomic DNA. Translation: EAW61132.1.
BC011677 mRNA. Translation: AAH11677.1.
BC031687 mRNA. Translation: AAH31687.1.
CCDSiCCDS34622.1. [Q9UJU6-2]
CCDS34623.1. [Q9UJU6-1]
CCDS47579.1. [Q9UJU6-3]
CCDS64633.1. [Q9UJU6-4]
CCDS64634.1. [Q9UJU6-5]
RefSeqiNP_001014436.1. NM_001014436.2. [Q9UJU6-1]
NP_001116428.1. NM_001122956.1. [Q9UJU6-3]
NP_001271242.1. NM_001284313.1. [Q9UJU6-5]
NP_001271244.1. NM_001284315.1. [Q9UJU6-4]
NP_054782.2. NM_014063.6. [Q9UJU6-2]
UniGeneiHs.436500.

3D structure databases

Select the link destinations:
PDBei
RCSB PDBi
PDBji
Links Updated
PDB entryMethodResolution (Å)ChainPositionsPDBsum
1X67NMR-A1-133[»]
ProteinModelPortaliQ9UJU6.
SMRiQ9UJU6.
ModBaseiSearch...
MobiDBiSearch...

Protein-protein interaction databases

BioGridi118809. 50 interactors.
IntActiQ9UJU6. 8 interactors.
MINTiMINT-1474845.
STRINGi9606.ENSP00000417653.

PTM databases

iPTMnetiQ9UJU6.
PhosphoSitePlusiQ9UJU6.

Polymorphism and mutation databases

BioMutaiDBNL.
DMDMi51316115.

2D gel databases

OGPiQ9UJU6.

Proteomic databases

EPDiQ9UJU6.
PaxDbiQ9UJU6.
PeptideAtlasiQ9UJU6.
PRIDEiQ9UJU6.
TopDownProteomicsiQ9UJU6-2. [Q9UJU6-2]

Protocols and materials databases

DNASUi28988.
Structural Biology KnowledgebaseSearch...

Genome annotation databases

EnsembliENST00000440166; ENSP00000415173; ENSG00000136279. [Q9UJU6-5]
ENST00000448521; ENSP00000411701; ENSG00000136279. [Q9UJU6-1]
ENST00000456905; ENSP00000416421; ENSG00000136279. [Q9UJU6-6]
ENST00000468694; ENSP00000417653; ENSG00000136279. [Q9UJU6-3]
ENST00000490734; ENSP00000417749; ENSG00000136279. [Q9UJU6-4]
ENST00000494774; ENSP00000419992; ENSG00000136279. [Q9UJU6-2]
GeneIDi28988.
KEGGihsa:28988.
UCSCiuc003tjo.5. human. [Q9UJU6-1]

Organism-specific databases

CTDi28988.
DisGeNETi28988.
GeneCardsiDBNL.
HGNCiHGNC:2696. DBNL.
HPAiHPA020265.
HPA027735.
MIMi610106. gene.
neXtProtiNX_Q9UJU6.
OpenTargetsiENSG00000136279.
PharmGKBiPA27164.
GenAtlasiSearch...

Phylogenomic databases

eggNOGiKOG3655. Eukaryota.
ENOG410XRVX. LUCA.
GeneTreeiENSGT00530000062953.
HOGENOMiHOG000008567.
HOVERGENiHBG051316.
InParanoidiQ9UJU6.
KOiK20520.
OMAiFQDTGPQ.
OrthoDBiEOG091G05KP.
PhylomeDBiQ9UJU6.
TreeFamiTF318935.

Enzyme and pathway databases

ReactomeiR-HSA-264870. Caspase-mediated cleavage of cytoskeletal proteins.
R-HSA-6794361. Interactions of neurexins and neuroligins at synapses.
R-HSA-6798695. Neutrophil degranulation.
SignaLinkiQ9UJU6.
SIGNORiQ9UJU6.

Miscellaneous databases

ChiTaRSiDBNL. human.
EvolutionaryTraceiQ9UJU6.
GeneWikiiDrebrin-like.
GenomeRNAii28988.
PMAP-CutDBQ9UJU6.
PROiQ9UJU6.
SOURCEiSearch...

Gene expression databases

BgeeiENSG00000136279.
CleanExiHS_DBNL.
ExpressionAtlasiQ9UJU6. baseline and differential.
GenevisibleiQ9UJU6. HS.

Family and domain databases

Gene3Di3.40.20.10. 1 hit.
InterProiIPR002108. ADF-H.
IPR029006. ADF-H/Gelsolin-like_dom.
IPR029923. Dbnl.
IPR001452. SH3_domain.
[Graphical view]
PANTHERiPTHR10829:SF12. PTHR10829:SF12. 1 hit.
PfamiPF00241. Cofilin_ADF. 1 hit.
PF14604. SH3_9. 1 hit.
[Graphical view]
SMARTiSM00102. ADF. 1 hit.
SM00326. SH3. 1 hit.
[Graphical view]
SUPFAMiSSF50044. SSF50044. 1 hit.
PROSITEiPS51263. ADF_H. 1 hit.
PS50002. SH3. 1 hit.
[Graphical view]
ProtoNetiSearch...

Entry informationi

Entry nameiDBNL_HUMAN
AccessioniPrimary (citable) accession number: Q9UJU6
Secondary accession number(s): A4D2I9
, B4DDP6, B4DEM2, C9J7P1, P84070, Q6IAI8, Q96F30, Q96K74, Q9HBN8, Q9NR72
Entry historyi
Integrated into UniProtKB/Swiss-Prot: August 16, 2004
Last sequence update: May 1, 2000
Last modified: November 30, 2016
This is version 154 of the entry and version 1 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

Keywords - Technical termi

3D-structure, Complete proteome, Reference proteome

Documents

  1. Human chromosome 7
    Human chromosome 7: entries, gene names and cross-references to MIM
  2. MIM cross-references
    Online Mendelian Inheritance in Man (MIM) cross-references in UniProtKB/Swiss-Prot
  3. PDB cross-references
    Index of Protein Data Bank (PDB) cross-references
  4. SIMILARITY comments
    Index of protein domains and families

Similar proteinsi

Links to similar proteins from the UniProt Reference Clusters (UniRef) at 100%, 90% and 50% sequence identity:
100%UniRef100 combines identical sequences and sub-fragments with 11 or more residues from any organism into one UniRef entry.
90%UniRef90 is built by clustering UniRef100 sequences that have at least 90% sequence identity to, and 80% overlap with, the longest sequence (a.k.a seed sequence).
50%UniRef50 is built by clustering UniRef90 seed sequences that have at least 50% sequence identity to, and 80% overlap with, the longest sequence in the cluster.