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Protein

Tyrosyl-DNA phosphodiesterase 1

Gene

TDP1

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

Functioni

DNA repair enzyme that can remove a variety of covalent adducts from DNA through hydrolysis of a 3'-phosphodiester bond, giving rise to DNA with a free 3' phosphate. Catalyzes the hydrolysis of dead-end complexes between DNA and the topoisomerase I active site tyrosine residue. Hydrolyzes 3'-phosphoglycolates on protruding 3' ends on DNA double-strand breaks due to DNA damage by radiation and free radicals. Acts on blunt-ended double-strand DNA breaks and on single-stranded DNA. Has low 3'exonuclease activity and can remove a single nucleoside from the 3'end of DNA and RNA molecules with 3'hydroxyl groups. Has no exonuclease activity towards DNA or RNA with a 3'phosphate.5 Publications

Kineticsi

kcat is 7 sec(-1) with single-stranded 5'-tyrosyl DNA as substrate.
  1. KM=0.08 µM for 14-mer single-stranded oligo with a 3'-phosphotyrosine1 Publication

    Sites

    Feature keyPosition(s)DescriptionActionsGraphical viewLength
    Active sitei263Nucleophile2 Publications1
    Binding sitei265Substrate1 Publication1
    Active sitei493Proton donor/acceptor2 Publications1
    Binding sitei495Substrate1 Publication1

    GO - Molecular functioni

    GO - Biological processi

    Keywordsi

    Molecular functionExonuclease, Hydrolase, Nuclease
    Biological processDNA damage, DNA repair

    Enzyme and pathway databases

    ReactomeiR-HSA-5693571 Nonhomologous End-Joining (NHEJ)
    SABIO-RKiQ9NUW8
    SIGNORiQ9NUW8

    Names & Taxonomyi

    Protein namesi
    Recommended name:
    Tyrosyl-DNA phosphodiesterase 1 (EC:3.1.4.-3 Publications)
    Short name:
    Tyr-DNA phosphodiesterase 1
    Gene namesi
    Name:TDP1
    OrganismiHomo sapiens (Human)
    Taxonomic identifieri9606 [NCBI]
    Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
    Proteomesi
    • UP000005640 Componenti: Chromosome 14

    Organism-specific databases

    EuPathDBiHostDB:ENSG00000042088.13
    HGNCiHGNC:18884 TDP1
    MIMi607198 gene
    neXtProtiNX_Q9NUW8

    Subcellular locationi

    Extracellular region or secreted Cytosol Plasma membrane Cytoskeleton Lysosome Endosome Peroxisome ER Golgi apparatus Nucleus Mitochondrion Manual annotation Automatic computational assertionGraphics by Christian Stolte; Source: COMPARTMENTS

    Keywords - Cellular componenti

    Cytoplasm, Nucleus

    Pathology & Biotechi

    Involvement in diseasei

    Spinocerebellar ataxia, autosomal recessive, with axonal neuropathy (SCAN1)5 Publications
    The disease is caused by mutations affecting the gene represented in this entry.
    Disease descriptionSpinocerebellar ataxia is a clinically and genetically heterogeneous group of cerebellar disorders. Patients show progressive incoordination of gait and often poor coordination of hands, speech and eye movements, due to degeneration of the cerebellum with variable involvement of the brainstem and spinal cord. SCAN1 is an autosomal recessive cerebellar ataxia (ARCA) associated with peripheral axonal motor and sensory neuropathy, distal muscular atrophy, pes cavus and steppage gait as seen in Charcot-Marie-Tooth neuropathy. All affected individuals have normal intelligence.
    See also OMIM:607250
    Feature keyPosition(s)DescriptionActionsGraphical viewLength
    Natural variantiVAR_017144493H → R in SCAN1; reduces enzyme activity and leads to the accumulation of covalent complexes between TDP1 and DNA. 5 PublicationsCorresponds to variant dbSNP:rs119467003EnsemblClinVar.1

    Mutagenesis

    Feature keyPosition(s)DescriptionActionsGraphical viewLength
    Mutagenesisi263H → A: Loss of activity. 2 Publications1
    Mutagenesisi265K → A: Abolishes hydrolysis of the covalent intermediate between the active site nucleophile and DNA. 2 Publications1
    Mutagenesisi265K → S: Reduces the activity to nearly undetectable levels. 2 Publications1
    Mutagenesisi283N → A: No effect. 1 Publication1
    Mutagenesisi294Q → A: Slightly reduced hydrolysis of the covalent intermediate between the active site nucleophile and DNA. 1 Publication1
    Mutagenesisi493H → A: 3000-fold reduction in activity; abolishes hydrolysis of the covalent intermediate between the active site nucleophile and DNA. 2 Publications1
    Mutagenesisi493H → N: 15000-fold reduction in activity. 2 Publications1
    Mutagenesisi495K → A: Abolishes hydrolysis of the covalent intermediate between the active site nucleophile and DNA. 2 Publications1
    Mutagenesisi495K → S: 125-fold reduction in activity. 2 Publications1
    Mutagenesisi516N → A: Reduced hydrolysis of the covalent intermediate between the active site nucleophile and DNA. 1 Publication1
    Mutagenesisi538E → A: Abolishes hydrolysis of the covalent intermediate between the active site nucleophile and DNA. 1 Publication1

    Keywords - Diseasei

    Disease mutation, Neurodegeneration

    Organism-specific databases

    DisGeNETi55775
    GeneReviewsiTDP1
    MalaCardsiTDP1
    MIMi607250 phenotype
    OpenTargetsiENSG00000042088
    Orphaneti94124 Spinocerebellar ataxia with axonal neuropathy type 1
    PharmGKBiPA421

    Chemistry databases

    ChEMBLiCHEMBL1075138

    Polymorphism and mutation databases

    BioMutaiTDP1
    DMDMi37999797

    PTM / Processingi

    Molecule processing

    Feature keyPosition(s)DescriptionActionsGraphical viewLength
    ChainiPRO_00002124861 – 608Tyrosyl-DNA phosphodiesterase 1Add BLAST608

    Amino acid modifications

    Feature keyPosition(s)DescriptionActionsGraphical viewLength
    Modified residuei61PhosphoserineCombined sources1
    Modified residuei147PhosphothreonineCombined sources1
    Modified residuei148PhosphoserineCombined sources1

    Post-translational modificationi

    Phosphorylated on serine and/or threonine residues, but not on tyrosine residues.1 Publication

    Keywords - PTMi

    Phosphoprotein

    Proteomic databases

    EPDiQ9NUW8
    MaxQBiQ9NUW8
    PaxDbiQ9NUW8
    PeptideAtlasiQ9NUW8
    PRIDEiQ9NUW8
    ProteomicsDBi82725

    PTM databases

    iPTMnetiQ9NUW8
    PhosphoSitePlusiQ9NUW8

    Expressioni

    Tissue specificityi

    Ubiquitously expressed. Similar expression throughout the central nervous system (whole brain, amygdala, caudate nucleus, cerebellum, cerebral cortex, frontal lobe, hippocampus, medulla oblongata, occipital lobe, putamen, substantia nigra, temporal lobe, thalamus, nucleus accumbens and spinal cord) and increased expression in testis and thymus.2 Publications

    Gene expression databases

    BgeeiENSG00000042088 Expressed in 170 organ(s), highest expression level in oocyte
    CleanExiHS_TDP1
    ExpressionAtlasiQ9NUW8 baseline and differential
    GenevisibleiQ9NUW8 HS

    Organism-specific databases

    HPAiHPA071317

    Interactioni

    Subunit structurei

    Monomer.1 Publication

    Sites

    Feature keyPosition(s)DescriptionActionsGraphical viewLength
    Sitei518Interaction with DNA1 Publication1

    Protein-protein interaction databases

    BioGridi120890, 12 interactors
    CORUMiQ9NUW8
    IntActiQ9NUW8, 5 interactors
    MINTiQ9NUW8
    STRINGi9606.ENSP00000337353

    Chemistry databases

    BindingDBiQ9NUW8

    Structurei

    Secondary structure

    1608
    Legend: HelixTurnBeta strandPDB Structure known for this area
    Show more details

    3D structure databases

    ProteinModelPortaliQ9NUW8
    SMRiQ9NUW8
    ModBaseiSearch...
    MobiDBiSearch...

    Miscellaneous databases

    EvolutionaryTraceiQ9NUW8

    Family & Domainsi

    Region

    Feature keyPosition(s)DescriptionActionsGraphical viewLength
    Regioni400 – 403Interaction with DNA1 Publication4

    Sequence similaritiesi

    Keywords - Domaini

    Repeat

    Phylogenomic databases

    eggNOGiKOG2031 Eukaryota
    ENOG410XQPZ LUCA
    GeneTreeiENSGT00390000002211
    HOGENOMiHOG000067775
    HOVERGENiHBG061242
    InParanoidiQ9NUW8
    KOiK10862
    OMAiQKFFSGS
    OrthoDBiEOG091G0DQF
    PhylomeDBiQ9NUW8
    TreeFamiTF105989

    Family and domain databases

    Gene3Di3.30.870.20, 1 hit
    InterProiView protein in InterPro
    IPR010347 Tdp1
    IPR027415 TDP_C
    PANTHERiPTHR12415 PTHR12415, 1 hit
    PfamiView protein in Pfam
    PF06087 Tyr-DNA_phospho, 1 hit

    Sequences (2+)i

    Sequence statusi: Complete.

    This entry describes 2 isoformsi produced by alternative splicing. AlignAdd to basket

    This entry has 2 described isoforms and 13 potential isoforms that are computationally mapped.Show allAlign All

    Isoform 1 (identifier: Q9NUW8-1) [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
    MSQEGDYGRW TISSSDESEE EKPKPDKPST SSLLCARQGA ANEPRYTCSE
    60 70 80 90 100
    AQKAAHKRKI SPVKFSNTDS VLPPKRQKSG SQEDLGWCLS SSDDELQPEM
    110 120 130 140 150
    PQKQAEKVVI KKEKDISAPN DGTAQRTENH GAPACHRLKE EEDEYETSGE
    160 170 180 190 200
    GQDIWDMLDK GNPFQFYLTR VSGVKPKYNS GALHIKDILS PLFGTLVSSA
    210 220 230 240 250
    QFNYCFDVDW LVKQYPPEFR KKPILLVHGD KREAKAHLHA QAKPYENISL
    260 270 280 290 300
    CQAKLDIAFG THHTKMMLLL YEEGLRVVIH TSNLIHADWH QKTQGIWLSP
    310 320 330 340 350
    LYPRIADGTH KSGESPTHFK ADLISYLMAY NAPSLKEWID VIHKHDLSET
    360 370 380 390 400
    NVYLIGSTPG RFQGSQKDNW GHFRLKKLLK DHASSMPNAE SWPVVGQFSS
    410 420 430 440 450
    VGSLGADESK WLCSEFKESM LTLGKESKTP GKSSVPLYLI YPSVENVRTS
    460 470 480 490 500
    LEGYPAGGSL PYSIQTAEKQ NWLHSYFHKW SAETSGRSNA MPHIKTYMRP
    510 520 530 540 550
    SPDFSKIAWF LVTSANLSKA AWGALEKNGT QLMIRSYELG VLFLPSAFGL
    560 570 580 590 600
    DSFKVKQKFF AGSQEPMATF PVPYDLPPEL YGSKDRPWIW NIPYVKAPDT

    HGNMWVPS
    Length:608
    Mass (Da):68,420
    Last modified:October 24, 2003 - v2
    Checksum:iF30202BD8329E5CE
    GO
    Isoform 2 (identifier: Q9NUW8-2) [UniParc]FASTAAdd to basket

    The sequence of this isoform differs from the canonical sequence as follows:
         1-25: MSQEGDYGRWTISSSDESEEEKPKP → MVISERLRLTSMPRPSLTRTSLSAR
         26-264: Missing.

    Note: No experimental confirmation available.
    Show »
    Length:369
    Mass (Da):41,630
    Checksum:i8674BAEED98FA149
    GO

    Computationally mapped potential isoform sequencesi

    There are 13 potential isoforms mapped to this entry.BLASTAlignShow allAdd to basket
    EntryEntry nameProtein names
    Gene namesLengthAnnotation
    G3V2F4G3V2F4_HUMAN
    Tyrosyl-DNA phosphodiesterase 1
    TDP1
    578Annotation score:
    E7EPD8E7EPD8_HUMAN
    Tyrosyl-DNA phosphodiesterase 1
    TDP1
    589Annotation score:
    G3V2U6G3V2U6_HUMAN
    Tyrosyl-DNA phosphodiesterase 1
    TDP1
    279Annotation score:
    G3V5B8G3V5B8_HUMAN
    Tyrosyl-DNA phosphodiesterase 1
    TDP1
    151Annotation score:
    G3V3Q0G3V3Q0_HUMAN
    Tyrosyl-DNA phosphodiesterase 1
    TDP1
    146Annotation score:
    G3V554G3V554_HUMAN
    Tyrosyl-DNA phosphodiesterase 1
    TDP1
    161Annotation score:
    G3V5F9G3V5F9_HUMAN
    Tyrosyl-DNA phosphodiesterase 1
    TDP1
    147Annotation score:
    H0YJ44H0YJ44_HUMAN
    Tyrosyl-DNA phosphodiesterase 1
    TDP1
    281Annotation score:
    G3V2J6G3V2J6_HUMAN
    Tyrosyl-DNA phosphodiesterase 1
    TDP1
    219Annotation score:
    G3V5H9G3V5H9_HUMAN
    Tyrosyl-DNA phosphodiesterase 1
    TDP1
    259Annotation score:
    There are more potential isoformsShow all

    Experimental Info

    Feature keyPosition(s)DescriptionActionsGraphical viewLength
    Sequence conflicti389A → P in BAA91997 (PubMed:14702039).Curated1
    Sequence conflicti511L → R in AAF65624 (PubMed:10521354).Curated1

    Natural variant

    Feature keyPosition(s)DescriptionActionsGraphical viewLength
    Natural variantiVAR_02581795E → D1 PublicationCorresponds to variant dbSNP:rs35114462EnsemblClinVar.1
    Natural variantiVAR_025818101P → L1 PublicationCorresponds to variant dbSNP:rs35455108EnsemblClinVar.1
    Natural variantiVAR_025819134A → T1 PublicationCorresponds to variant dbSNP:rs28365054EnsemblClinVar.1
    Natural variantiVAR_025820187D → G1 PublicationCorresponds to variant dbSNP:rs35271143Ensembl.1
    Natural variantiVAR_025821304R → Q1 PublicationCorresponds to variant dbSNP:rs34452707EnsemblClinVar.1
    Natural variantiVAR_017144493H → R in SCAN1; reduces enzyme activity and leads to the accumulation of covalent complexes between TDP1 and DNA. 5 PublicationsCorresponds to variant dbSNP:rs119467003EnsemblClinVar.1
    Natural variantiVAR_017145566P → L in autosomal recessive or sporadic spinocerebellar ataxia affected Japanese individuals. 1 PublicationCorresponds to variant dbSNP:rs767298655Ensembl.1
    Natural variantiVAR_025822569T → A1 PublicationCorresponds to variant dbSNP:rs35973343EnsemblClinVar.1

    Alternative sequence

    Feature keyPosition(s)DescriptionActionsGraphical viewLength
    Alternative sequenceiVSP_0557651 – 25MSQEG…EKPKP → MVISERLRLTSMPRPSLTRT SLSAR in isoform 2. 1 PublicationAdd BLAST25
    Alternative sequenceiVSP_05576626 – 264Missing in isoform 2. 1 PublicationAdd BLAST239

    Sequence databases

    Select the link destinations:
    EMBLi
    GenBanki
    DDBJi
    Links Updated
    BX161451 mRNA Translation: CAD61915.1
    AK001952 mRNA Translation: BAA91997.1
    DQ367843 Genomic DNA Translation: ABC79301.1
    AL137128 Genomic DNA No translation available.
    BC015474 mRNA Translation: AAH15474.1
    AF182002 mRNA Translation: AAF65623.1
    AF182003 mRNA Translation: AAF65624.1
    CCDSiCCDS9888.1 [Q9NUW8-1]
    RefSeqiNP_001008744.1, NM_001008744.1 [Q9NUW8-1]
    NP_060789.2, NM_018319.3 [Q9NUW8-1]
    XP_005267904.1, XM_005267847.2
    XP_005267905.1, XM_005267848.2 [Q9NUW8-1]
    XP_006720260.1, XM_006720197.3 [Q9NUW8-1]
    XP_006720261.1, XM_006720198.3
    XP_011535244.1, XM_011536942.2 [Q9NUW8-1]
    XP_011535245.1, XM_011536943.2
    XP_016876928.1, XM_017021439.1 [Q9NUW8-1]
    XP_016876931.1, XM_017021442.1
    XP_016876932.1, XM_017021443.1
    XP_016876933.1, XM_017021444.1
    UniGeneiHs.209945

    Genome annotation databases

    EnsembliENST00000335725; ENSP00000337353; ENSG00000042088 [Q9NUW8-1]
    ENST00000393454; ENSP00000377099; ENSG00000042088 [Q9NUW8-1]
    GeneIDi55775
    KEGGihsa:55775
    UCSCiuc001xxy.4 human [Q9NUW8-1]

    Keywords - Coding sequence diversityi

    Alternative splicing, Polymorphism

    Similar proteinsi

    Cross-referencesi

    Web resourcesi

    NIEHS-SNPs

    Sequence databases

    Select the link destinations:
    EMBLi
    GenBanki
    DDBJi
    Links Updated
    BX161451 mRNA Translation: CAD61915.1
    AK001952 mRNA Translation: BAA91997.1
    DQ367843 Genomic DNA Translation: ABC79301.1
    AL137128 Genomic DNA No translation available.
    BC015474 mRNA Translation: AAH15474.1
    AF182002 mRNA Translation: AAF65623.1
    AF182003 mRNA Translation: AAF65624.1
    CCDSiCCDS9888.1 [Q9NUW8-1]
    RefSeqiNP_001008744.1, NM_001008744.1 [Q9NUW8-1]
    NP_060789.2, NM_018319.3 [Q9NUW8-1]
    XP_005267904.1, XM_005267847.2
    XP_005267905.1, XM_005267848.2 [Q9NUW8-1]
    XP_006720260.1, XM_006720197.3 [Q9NUW8-1]
    XP_006720261.1, XM_006720198.3
    XP_011535244.1, XM_011536942.2 [Q9NUW8-1]
    XP_011535245.1, XM_011536943.2
    XP_016876928.1, XM_017021439.1 [Q9NUW8-1]
    XP_016876931.1, XM_017021442.1
    XP_016876932.1, XM_017021443.1
    XP_016876933.1, XM_017021444.1
    UniGeneiHs.209945

    3D structure databases

    Select the link destinations:
    PDBei
    RCSB PDBi
    PDBji
    Links Updated
    PDB entryMethodResolution (Å)ChainPositionsPDBsum
    1JY1X-ray1.69A149-608[»]
    1MU7X-ray2.00A/B149-608[»]
    1MU9X-ray2.05A/B149-608[»]
    1NOPX-ray2.30A/B149-608[»]
    1QZQX-ray2.40A/B149-608[»]
    1RFFX-ray1.70A/B149-608[»]
    1RFIX-ray2.20A/B149-608[»]
    1RG1X-ray2.10A/B149-608[»]
    1RG2X-ray2.10A/B149-608[»]
    1RGTX-ray2.00A/B149-608[»]
    1RGUX-ray2.22A/B149-608[»]
    1RH0X-ray2.30A/B149-608[»]
    5NW9X-ray2.04A/B149-608[»]
    5NWAX-ray3.20A/B149-608[»]
    ProteinModelPortaliQ9NUW8
    SMRiQ9NUW8
    ModBaseiSearch...
    MobiDBiSearch...

    Protein-protein interaction databases

    BioGridi120890, 12 interactors
    CORUMiQ9NUW8
    IntActiQ9NUW8, 5 interactors
    MINTiQ9NUW8
    STRINGi9606.ENSP00000337353

    Chemistry databases

    BindingDBiQ9NUW8
    ChEMBLiCHEMBL1075138

    PTM databases

    iPTMnetiQ9NUW8
    PhosphoSitePlusiQ9NUW8

    Polymorphism and mutation databases

    BioMutaiTDP1
    DMDMi37999797

    Proteomic databases

    EPDiQ9NUW8
    MaxQBiQ9NUW8
    PaxDbiQ9NUW8
    PeptideAtlasiQ9NUW8
    PRIDEiQ9NUW8
    ProteomicsDBi82725

    Protocols and materials databases

    Structural Biology KnowledgebaseSearch...

    Genome annotation databases

    EnsembliENST00000335725; ENSP00000337353; ENSG00000042088 [Q9NUW8-1]
    ENST00000393454; ENSP00000377099; ENSG00000042088 [Q9NUW8-1]
    GeneIDi55775
    KEGGihsa:55775
    UCSCiuc001xxy.4 human [Q9NUW8-1]

    Organism-specific databases

    CTDi55775
    DisGeNETi55775
    EuPathDBiHostDB:ENSG00000042088.13
    GeneCardsiTDP1
    GeneReviewsiTDP1
    HGNCiHGNC:18884 TDP1
    HPAiHPA071317
    MalaCardsiTDP1
    MIMi607198 gene
    607250 phenotype
    neXtProtiNX_Q9NUW8
    OpenTargetsiENSG00000042088
    Orphaneti94124 Spinocerebellar ataxia with axonal neuropathy type 1
    PharmGKBiPA421
    GenAtlasiSearch...

    Phylogenomic databases

    eggNOGiKOG2031 Eukaryota
    ENOG410XQPZ LUCA
    GeneTreeiENSGT00390000002211
    HOGENOMiHOG000067775
    HOVERGENiHBG061242
    InParanoidiQ9NUW8
    KOiK10862
    OMAiQKFFSGS
    OrthoDBiEOG091G0DQF
    PhylomeDBiQ9NUW8
    TreeFamiTF105989

    Enzyme and pathway databases

    ReactomeiR-HSA-5693571 Nonhomologous End-Joining (NHEJ)
    SABIO-RKiQ9NUW8
    SIGNORiQ9NUW8

    Miscellaneous databases

    EvolutionaryTraceiQ9NUW8
    GeneWikiiTDP1
    GenomeRNAii55775
    PROiPR:Q9NUW8
    SOURCEiSearch...

    Gene expression databases

    BgeeiENSG00000042088 Expressed in 170 organ(s), highest expression level in oocyte
    CleanExiHS_TDP1
    ExpressionAtlasiQ9NUW8 baseline and differential
    GenevisibleiQ9NUW8 HS

    Family and domain databases

    Gene3Di3.30.870.20, 1 hit
    InterProiView protein in InterPro
    IPR010347 Tdp1
    IPR027415 TDP_C
    PANTHERiPTHR12415 PTHR12415, 1 hit
    PfamiView protein in Pfam
    PF06087 Tyr-DNA_phospho, 1 hit
    ProtoNetiSearch...

    Entry informationi

    Entry nameiTYDP1_HUMAN
    AccessioniPrimary (citable) accession number: Q9NUW8
    Secondary accession number(s): Q2HXX4
    , Q86TV8, Q96BK7, Q9NZM7, Q9NZM8
    Entry historyiIntegrated into UniProtKB/Swiss-Prot: October 24, 2003
    Last sequence update: October 24, 2003
    Last modified: November 7, 2018
    This is version 153 of the entry and version 2 of the sequence. See 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. SIMILARITY comments
      Index of protein domains and families
    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. PDB cross-references
      Index of Protein Data Bank (PDB) cross-references
    6. Human chromosome 14
      Human chromosome 14: entries, gene names and cross-references to MIM
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