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Protein

Mitogen-activated protein kinase 7

Gene

Mapk7

Organism
Mus musculus (Mouse)
Status
Reviewed-Annotation score: Annotation score: 5 out of 5-Experimental evidence at protein leveli

Functioni

Plays a role in various cellular processes such as proliferation, differentiation and cell survival. The upstream activator of MAPK7 is the MAPK kinase MAP2K5. Upon activation, it translocates to the nucleus and phosphorylates various downstream targets including MEF2C. EGF activates MAPK7 through a Ras-independent and MAP2K5-dependent pathway. May have a role in muscle cell differentiation. May be important for endothelial function and maintenance of blood vessel integrity. MAP2K5 and MAPK7 interact specifically with one another and not with MEK1/ERK1 or MEK2/ERK2 pathways. Phosphorylates SGK1 at Ser-78 and this is required for growth factor-induced cell cycle progression (By similarity). Involved in the regulation of p53/TP53 by disrupting the PML-MDM2 interaction (By similarity).By similarity5 Publications

Catalytic activityi

ATP + a protein = ADP + a phosphoprotein.

Cofactori

Mg2+By similarity

Enzyme regulationi

Activated by tyrosine and threonine phosphorylation. Activated in response to hyperosmolarity, hydrogen peroxide, and epidermal growth factor (EGF) (By similarity).By similarity

Sites

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Binding sitei84ATPPROSITE-ProRule annotation1
Active sitei182Proton acceptorPROSITE-ProRule annotation1

Regions

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Nucleotide bindingi61 – 69ATPPROSITE-ProRule annotation9

GO - Molecular functioni

  • ATP binding Source: UniProtKB-KW
  • mitogen-activated protein kinase binding Source: MGI
  • protein kinase activity Source: UniProtKB
  • protein serine/threonine kinase activity Source: MGI

GO - Biological processi

Complete GO annotation...

Keywords - Molecular functioni

Kinase, Serine/threonine-protein kinase, Transferase

Keywords - Biological processi

Cell cycle, Differentiation

Keywords - Ligandi

ATP-binding, Nucleotide-binding

Enzyme and pathway databases

ReactomeiR-MMU-198753. ERK/MAPK targets.
R-MMU-198765. Signalling to ERK5.
R-MMU-202670. ERKs are inactivated.
R-MMU-2559582. Senescence-Associated Secretory Phenotype (SASP).
R-MMU-881907. Gastrin-CREB signalling pathway via PKC and MAPK.
R-MMU-8853659. RET signaling.

Names & Taxonomyi

Protein namesi
Recommended name:
Mitogen-activated protein kinase 7 (EC:2.7.11.24)
Short name:
MAP kinase 7
Short name:
MAPK 7
Alternative name(s):
Big MAP kinase 1
Short name:
BMK-1
Extracellular signal-regulated kinase 5
Short name:
ERK-5
Gene namesi
Name:Mapk7
Synonyms:Bmk1, Erk5
OrganismiMus musculus (Mouse)
Taxonomic identifieri10090 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresGliresRodentiaSciurognathiMuroideaMuridaeMurinaeMusMus
Proteomesi
  • UP000000589 Componenti: Chromosome 11

Organism-specific databases

MGIiMGI:1346347. Mapk7.

Subcellular locationi

  • Cytoplasm 1 Publication
  • Nucleus 1 Publication
  • NucleusPML body By similarity

  • Note: Translocates to the nucleus upon activation.By similarity
Isoform 1 :
  • Cytoplasm By similarity
  • Nucleus By similarity

  • Note: Isoform 1 is detected in cytoplasm and nucleus.By similarity
Isoform 2 :
  • Nucleus By similarity

  • Note: Isoform 2 is detected only in the nucleus. Translocates to the nucleus upon activation (By similarity).By similarity
Isoform 3 :
  • Nucleus By similarity

  • Note: Isoform 2 is detected only in the nucleus. Translocates to the nucleus upon activation (By similarity).By similarity

GO - Cellular componenti

  • cytoplasm Source: UniProtKB
  • cytosol Source: MGI
  • nucleoplasm Source: MGI
  • nucleus Source: MGI
  • PML body Source: UniProtKB
Complete GO annotation...

Keywords - Cellular componenti

Cytoplasm, Nucleus

PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Initiator methionineiRemovedBy similarity
ChainiPRO_00001862612 – 806Mitogen-activated protein kinase 7Add BLAST805

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Modified residuei2N-acetylalanineBy similarity1
Modified residuei710PhosphoserineBy similarity1
Modified residuei723PhosphothreonineBy similarity1

Post-translational modificationi

Dually phosphorylated on Thr-219 and Tyr-221, which activates the enzyme.By similarity

Keywords - PTMi

Acetylation, Phosphoprotein

Proteomic databases

PaxDbiQ9WVS8.
PeptideAtlasiQ9WVS8.
PRIDEiQ9WVS8.

PTM databases

iPTMnetiQ9WVS8.
PhosphoSitePlusiQ9WVS8.

Expressioni

Tissue specificityi

Detected in testis, brain, kidney, lung and heart. Detected in total embryo (at protein level).1 Publication

Gene expression databases

BgeeiENSMUSG00000001034.
CleanExiMM_MAPK7.
ExpressionAtlasiQ9WVS8. baseline and differential.
GenevisibleiQ9WVS8. MM.

Interactioni

Subunit structurei

Interacts with MAP2K5 (By similarity). Forms oligomers. Interacts with MEF2A, MEF2C and MEF2D; the interaction phosphorylates the MEF2s and enhances transcriptional activity of MEF2A, MEF2C but not MEF2D (By similarity). Interacts with SGK1 (By similarity). Interacts with PML (By similarity).By similarity

GO - Molecular functioni

Protein-protein interaction databases

BioGridi204806. 4 interactors.
STRINGi10090.ENSMUSP00000078087.

Structurei

3D structure databases

ProteinModelPortaliQ9WVS8.
SMRiQ9WVS8.
ModBaseiSearch...
MobiDBiSearch...

Family & Domainsi

Domains and Repeats

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Domaini55 – 347Protein kinasePROSITE-ProRule annotationAdd BLAST293

Region

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Regioni2 – 77Required for cytoplasmic targetingAdd BLAST76
Regioni78 – 139Required for binding to MAP2K5Add BLAST62
Regioni140 – 406Necessary for oligomerizationAdd BLAST267
Regioni407 – 806May not be required for kinase activity; required to stimulate MEF2C activityAdd BLAST400

Motif

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Motifi219 – 221TXY3
Motifi505 – 539Nuclear localization signalAdd BLAST35

Compositional bias

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Compositional biasi434 – 465Pro-richAdd BLAST32
Compositional biasi521 – 524Poly-Arg4
Compositional biasi578 – 700Pro-richAdd BLAST123

Domaini

The second proline-rich region may interact with actin targeting the kinase to a specific location in the cell.
The TXY motif contains the threonine and tyrosine residues whose phosphorylation activates the MAP kinases.

Sequence similaritiesi

Contains 1 protein kinase domain.PROSITE-ProRule annotation

Phylogenomic databases

eggNOGiKOG0660. Eukaryota.
ENOG410XNY0. LUCA.
GeneTreeiENSGT00550000074298.
HOVERGENiHBG108137.
InParanoidiQ9WVS8.
KOiK04464.
OMAiIQDFHKK.
OrthoDBiEOG091G08QL.
PhylomeDBiQ9WVS8.
TreeFamiTF105099.

Family and domain databases

InterProiIPR011009. Kinase-like_dom.
IPR003527. MAP_kinase_CS.
IPR000719. Prot_kinase_dom.
IPR017441. Protein_kinase_ATP_BS.
IPR008271. Ser/Thr_kinase_AS.
[Graphical view]
PfamiPF00069. Pkinase. 1 hit.
[Graphical view]
SMARTiSM00220. S_TKc. 1 hit.
[Graphical view]
SUPFAMiSSF56112. SSF56112. 1 hit.
PROSITEiPS01351. MAPK. 1 hit.
PS00107. PROTEIN_KINASE_ATP. 1 hit.
PS50011. PROTEIN_KINASE_DOM. 1 hit.
PS00108. PROTEIN_KINASE_ST. 1 hit.
[Graphical view]

Sequences (5)i

Sequence statusi: Complete.

Sequence processingi: The displayed sequence is further processed into a mature form.

This entry describes 5 isoformsi produced by alternative splicing. AlignAdd to basket

Isoform 1 (identifier: Q9WVS8-1) [UniParc]FASTAAdd to basket
Also known as: Big MAP kinase 1a, mERK5a

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
MAEPLKEEDG EDGSGEPPGR VKAEPVHTAA SVVAKNLALL KARSFDVTFD
60 70 80 90 100
VGDEYEIIET IGNGAYGVVS SARRRLTGQQ VAIKKIPNAF DVVTNAKRTL
110 120 130 140 150
RELKILKHFK HDNIIAIKDI LKPTVPYGEF RSVYVVLDLM ESDLHQIIHS
160 170 180 190 200
SQPLTLEHVR YFLYQLLRGL KYMHSAQVIH RDLKPSNLLV NENCELKIGD
210 220 230 240 250
FGMARGLCTS PAEHQYFMTE YVATRWYRAP ELMLSLHEYT QAIDLWSVGC
260 270 280 290 300
IFGEMLARRQ LFPGKNYVHQ LQLIMMVLGT PSPAVIQAVG AERVRAYIQS
310 320 330 340 350
LPPRQPVPWE TVYPGADRQA LSLLGRMLRF EPSARISAAA ALRHPFLAKY
360 370 380 390 400
HDPDDEPDCA PPFDFAFDRE ALTRERIKEA IVAEIEDFHA RREGIRQQIR
410 420 430 440 450
FQPSLQPVAS EPVCPDVEMP SPWAPSGDCA MESPPPALPP CSDPAPDTVD
460 470 480 490 500
LTLQPAPPAS ELAPPKREGA ISDNTKAALK AALLKSLRSR LRDGPSAPLE
510 520 530 540 550
APEPRKPVTA QERQREREEK RRRRQERAKE REKRRQERER KERGAGTLGG
560 570 580 590 600
PSTDPLAGLV LSDNDRSLLE RWTRMARPPA PAPAPAPAPA PAPSSAQPTS
610 620 630 640 650
TPTGPVSQST GPLQPAGSIP GPASQPVCPP PGPVPQPAGP IPAPLQTAPS
660 670 680 690 700
TSLLASQSLV PPSGLPGSGA PEVLPYFPSG PPPPDPGLTP QPSTSESPDV
710 720 730 740 750
NLVTQQLSKS QVEDPLPPVF SGTPKGSGAG YGVGFDLEEF LNQSFDMGVA
760 770 780 790 800
DGPQDGQADS ASLSASLLAD WLEGHGMNPA DIESLQREIQ MDSPMLLSDL

PDLQEP
Length:806
Mass (Da):87,733
Last modified:November 1, 1999 - v1
Checksum:iE7CC41C4BBDE0633
GO
Isoform 2 (identifier: Q9WVS8-2) [UniParc]FASTAAdd to basket
Also known as: Big MAP kinase 1b, mERK5b

The sequence of this isoform differs from the canonical sequence as follows:
     1-77: MAEPLKEEDG...VVSSARRRLT → MCGLLSRG

Note: May not have a kinase activity. May not stimulate MEF2C activity. May function as dominant negative inhibitor of the ERK5 signaling pathway.
Show »
Length:737
Mass (Da):80,380
Checksum:i29CEDCA390068527
GO
Isoform 3 (identifier: Q9WVS8-3) [UniParc]FASTAAdd to basket
Also known as: Big MAP kinase 1c, mERK5c

The sequence of this isoform differs from the canonical sequence as follows:
     1-139: Missing.

Note: May not have a kinase activity. May not stimulate MEF2C activity. May function as dominant negative inhibitor of the ERK5 signaling pathway.
Show »
Length:667
Mass (Da):72,463
Checksum:i00C42D9700759F7E
GO
Isoform 4 (identifier: Q9WVS8-4) [UniParc]FASTAAdd to basket
Also known as: Big MAP kinase 1d, mERK5-T

The sequence of this isoform differs from the canonical sequence as follows:
     493-806: DGPSAPLEAP...LSDLPDLQEP → GGVWAQQLSG

Note: Unable to translocate from the cytoplasm to the nucleus.
Show »
Length:502
Mass (Da):55,914
Checksum:i905BD56BC58913D8
GO
Isoform 5 (identifier: Q9WVS8-5) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     757-806: Missing.

Note: No experimental confirmation available.
Show »
Length:756
Mass (Da):82,314
Checksum:iC0CCFA5D2F8C84F5
GO

Experimental Info

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Sequence conflicti191N → D in BAE28357 (PubMed:16141072).Curated1
Sequence conflicti308P → H in BAE28357 (PubMed:16141072).Curated1
Sequence conflicti331E → K in AAI00399 (PubMed:15489334).Curated1
Sequence conflicti409A → V in AAD39394 (PubMed:11139578).Curated1
Sequence conflicti409A → V in AAD39395 (PubMed:11139578).Curated1
Sequence conflicti409A → V in AAD39396 (PubMed:11139578).Curated1
Sequence conflicti424A → R in AAD39394 (PubMed:11139578).Curated1
Sequence conflicti424A → R in AAD39395 (PubMed:11139578).Curated1
Sequence conflicti424A → R in AAD39396 (PubMed:11139578).Curated1
Sequence conflicti578P → PPAPA in BAE28357 (PubMed:16141072).Curated1
Sequence conflicti578P → PPAPA in BAE33103 (PubMed:16141072).Curated1
Sequence conflicti588A → R in AAD39394 (PubMed:11139578).Curated1
Sequence conflicti588A → R in AAD39395 (PubMed:11139578).Curated1
Sequence conflicti588A → R in AAD39396 (PubMed:11139578).Curated1
Sequence conflicti592A → Q in AAD39394 (PubMed:11139578).Curated1
Sequence conflicti592A → Q in AAD39395 (PubMed:11139578).Curated1
Sequence conflicti592A → Q in AAD39396 (PubMed:11139578).Curated1
Sequence conflicti618S → C in AAD39394 (PubMed:11139578).Curated1
Sequence conflicti618S → C in AAD39395 (PubMed:11139578).Curated1
Sequence conflicti618S → C in AAD39396 (PubMed:11139578).Curated1
Sequence conflicti642P → A in AAD39394 (PubMed:11139578).Curated1
Sequence conflicti642P → A in AAD39395 (PubMed:11139578).Curated1
Sequence conflicti642P → A in AAD39396 (PubMed:11139578).Curated1
Sequence conflicti803L → V in BAE28357 (PubMed:16141072).Curated1

Alternative sequence

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Alternative sequenceiVSP_0352011 – 139Missing in isoform 3. 1 PublicationAdd BLAST139
Alternative sequenceiVSP_0352021 – 77MAEPL…RRRLT → MCGLLSRG in isoform 2. 2 PublicationsAdd BLAST77
Alternative sequenceiVSP_035203493 – 806DGPSA…DLQEP → GGVWAQQLSG in isoform 4. 1 PublicationAdd BLAST314
Alternative sequenceiVSP_035204757 – 806Missing in isoform 5. 1 PublicationAdd BLAST50

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AB019373 mRNA. Translation: BAA82039.1.
AF126159 mRNA. Translation: AAD39394.1.
AF126160 mRNA. Translation: AAD39395.1.
AF126161 mRNA. Translation: AAD39396.1.
AK148119 mRNA. Translation: BAE28357.1.
AK155187 mRNA. Translation: BAE33103.1.
AY534740 mRNA. Translation: AAS38576.1.
BC100398 mRNA. Translation: AAI00399.1.
AL604029 Genomic DNA. Translation: CAI24182.1.
AL604029 Genomic DNA. Translation: CAI24184.1.
AL604029 Genomic DNA. Translation: CAI24185.1.
CCDSiCCDS24814.1. [Q9WVS8-1]
CCDS70205.1. [Q9WVS8-2]
RefSeqiNP_001277962.1. NM_001291033.1. [Q9WVS8-2]
NP_001277963.1. NM_001291034.1. [Q9WVS8-1]
NP_001277964.1. NM_001291035.1. [Q9WVS8-3]
NP_001277965.1. NM_001291036.1. [Q9WVS8-3]
NP_001277966.1. NM_001291037.1.
NP_035971.1. NM_011841.2. [Q9WVS8-1]
XP_006533340.1. XM_006533277.3. [Q9WVS8-1]
XP_006533343.1. XM_006533280.3. [Q9WVS8-3]
XP_006533344.1. XM_006533281.3. [Q9WVS8-3]
XP_006533347.1. XM_006533284.3. [Q9WVS8-3]
XP_006533348.1. XM_006533285.3. [Q9WVS8-3]
XP_006533349.1. XM_006533286.3. [Q9WVS8-3]
UniGeneiMm.38172.

Genome annotation databases

EnsembliENSMUST00000079080; ENSMUSP00000078087; ENSMUSG00000001034. [Q9WVS8-1]
ENSMUST00000108714; ENSMUSP00000104354; ENSMUSG00000001034. [Q9WVS8-2]
GeneIDi23939.
KEGGimmu:23939.
UCSCiuc007jho.2. mouse. [Q9WVS8-2]
uc007jhp.2. mouse. [Q9WVS8-1]
uc007jht.1. mouse. [Q9WVS8-4]

Keywords - Coding sequence diversityi

Alternative splicing

Cross-referencesi

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AB019373 mRNA. Translation: BAA82039.1.
AF126159 mRNA. Translation: AAD39394.1.
AF126160 mRNA. Translation: AAD39395.1.
AF126161 mRNA. Translation: AAD39396.1.
AK148119 mRNA. Translation: BAE28357.1.
AK155187 mRNA. Translation: BAE33103.1.
AY534740 mRNA. Translation: AAS38576.1.
BC100398 mRNA. Translation: AAI00399.1.
AL604029 Genomic DNA. Translation: CAI24182.1.
AL604029 Genomic DNA. Translation: CAI24184.1.
AL604029 Genomic DNA. Translation: CAI24185.1.
CCDSiCCDS24814.1. [Q9WVS8-1]
CCDS70205.1. [Q9WVS8-2]
RefSeqiNP_001277962.1. NM_001291033.1. [Q9WVS8-2]
NP_001277963.1. NM_001291034.1. [Q9WVS8-1]
NP_001277964.1. NM_001291035.1. [Q9WVS8-3]
NP_001277965.1. NM_001291036.1. [Q9WVS8-3]
NP_001277966.1. NM_001291037.1.
NP_035971.1. NM_011841.2. [Q9WVS8-1]
XP_006533340.1. XM_006533277.3. [Q9WVS8-1]
XP_006533343.1. XM_006533280.3. [Q9WVS8-3]
XP_006533344.1. XM_006533281.3. [Q9WVS8-3]
XP_006533347.1. XM_006533284.3. [Q9WVS8-3]
XP_006533348.1. XM_006533285.3. [Q9WVS8-3]
XP_006533349.1. XM_006533286.3. [Q9WVS8-3]
UniGeneiMm.38172.

3D structure databases

ProteinModelPortaliQ9WVS8.
SMRiQ9WVS8.
ModBaseiSearch...
MobiDBiSearch...

Protein-protein interaction databases

BioGridi204806. 4 interactors.
STRINGi10090.ENSMUSP00000078087.

PTM databases

iPTMnetiQ9WVS8.
PhosphoSitePlusiQ9WVS8.

Proteomic databases

PaxDbiQ9WVS8.
PeptideAtlasiQ9WVS8.
PRIDEiQ9WVS8.

Protocols and materials databases

Structural Biology KnowledgebaseSearch...

Genome annotation databases

EnsembliENSMUST00000079080; ENSMUSP00000078087; ENSMUSG00000001034. [Q9WVS8-1]
ENSMUST00000108714; ENSMUSP00000104354; ENSMUSG00000001034. [Q9WVS8-2]
GeneIDi23939.
KEGGimmu:23939.
UCSCiuc007jho.2. mouse. [Q9WVS8-2]
uc007jhp.2. mouse. [Q9WVS8-1]
uc007jht.1. mouse. [Q9WVS8-4]

Organism-specific databases

CTDi5598.
MGIiMGI:1346347. Mapk7.

Phylogenomic databases

eggNOGiKOG0660. Eukaryota.
ENOG410XNY0. LUCA.
GeneTreeiENSGT00550000074298.
HOVERGENiHBG108137.
InParanoidiQ9WVS8.
KOiK04464.
OMAiIQDFHKK.
OrthoDBiEOG091G08QL.
PhylomeDBiQ9WVS8.
TreeFamiTF105099.

Enzyme and pathway databases

ReactomeiR-MMU-198753. ERK/MAPK targets.
R-MMU-198765. Signalling to ERK5.
R-MMU-202670. ERKs are inactivated.
R-MMU-2559582. Senescence-Associated Secretory Phenotype (SASP).
R-MMU-881907. Gastrin-CREB signalling pathway via PKC and MAPK.
R-MMU-8853659. RET signaling.

Miscellaneous databases

PROiQ9WVS8.
SOURCEiSearch...

Gene expression databases

BgeeiENSMUSG00000001034.
CleanExiMM_MAPK7.
ExpressionAtlasiQ9WVS8. baseline and differential.
GenevisibleiQ9WVS8. MM.

Family and domain databases

InterProiIPR011009. Kinase-like_dom.
IPR003527. MAP_kinase_CS.
IPR000719. Prot_kinase_dom.
IPR017441. Protein_kinase_ATP_BS.
IPR008271. Ser/Thr_kinase_AS.
[Graphical view]
PfamiPF00069. Pkinase. 1 hit.
[Graphical view]
SMARTiSM00220. S_TKc. 1 hit.
[Graphical view]
SUPFAMiSSF56112. SSF56112. 1 hit.
PROSITEiPS01351. MAPK. 1 hit.
PS00107. PROTEIN_KINASE_ATP. 1 hit.
PS50011. PROTEIN_KINASE_DOM. 1 hit.
PS00108. PROTEIN_KINASE_ST. 1 hit.
[Graphical view]
ProtoNetiSearch...

Entry informationi

Entry nameiMK07_MOUSE
AccessioniPrimary (citable) accession number: Q9WVS8
Secondary accession number(s): Q3U2N7
, Q3UG52, Q497T0, Q5NCN6, Q5NCN7, Q5NCN9, Q6QLU8, Q9R1D9, Q9WVF3, Q9WVF4
Entry historyi
Integrated into UniProtKB/Swiss-Prot: February 21, 2001
Last sequence update: November 1, 1999
Last modified: November 2, 2016
This is version 146 of the entry and version 1 of the sequence. [Complete history]
Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programChordata Protein Annotation Program

Miscellaneousi

Keywords - Technical termi

Complete proteome, Reference proteome

Documents

  1. MGD cross-references
    Mouse Genome Database (MGD) cross-references in UniProtKB/Swiss-Prot
  2. Human and mouse protein kinases
    Human and mouse protein kinases: classification and index
  3. 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.