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TP53 Gene

protein-coding   GIFtS: 80
GCID: GC17M007565

Tumor Protein P53

  See TP53-related diseases
at  

(According to 1HGNC, 2Entrez Gene,
3UniProtKB/Swiss-Prot, 4UniProtKB/TrEMBL, 5OMIM, 6GeneLoc, 7Ensembl, 8DME, 9miRBase, 10fRNAdb, 12H-InvDB, 13NCBI, 14NONCODE, and/or 15RNAdb)
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Aliases
Tumor Protein P531 2     TRP532
P532 3 5     Cellular Tumor Antigen P532
Antigen NY-CO-132 3     Mutant Tumor Protein 532
Phosphoprotein P532 3     P53 Tumor Suppressor2
BCC72 5     Transformation-Related Protein 532
LFS12 5     Tumor Protein 532
Li-Fraumeni Syndrome1     Tumor Suppressor P533

External Ids:    HGNC: 119981   Entrez Gene: 71572   Ensembl: ENSG000001415107   OMIM: 1911705   UniProtKB: P046373   

Export aliases for TP53 gene to outside databases

Previous GC identifers: GC17P008026 GC17M008311 GC17M007514 GC17M007772 GC17M007512 GC17M007465


(According to Entrez Gene, GeneCards, Tocris Bioscience, Wikipedia's Gene Wiki, PharmGKB,
UniProtKB/Swiss-Prot, and/or UniProtKB/TrEMBL)
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Entrez Gene summary for TP53 Gene:
This gene encodes a tumor suppressor protein containing transcriptional activation, DNA binding, and
oligomerization domains. The encoded protein responds to diverse cellular stresses to regulate expression of
target genes, thereby inducing cell cycle arrest, apoptosis, senescence, DNA repair, or changes in metabolism.
Mutations in this gene are associated with a variety of human cancers, including hereditary cancers such as
Li-Fraumeni syndrome. Alternative splicing of this gene and the use of alternate promoters result in multiple
transcript variants and isoforms. Additional isoforms have also been shown to result from the use of alternate
translation initiation codons (PMIDs: 12032546, 20937277). (provided by RefSeq, Feb 2013)

GeneCards Summary for TP53 Gene:
TP53 (tumor protein p53) is a protein-coding gene. Diseases associated with TP53 include hepatocellular carcinoma, and osteosarcoma. GO annotations related to this gene include protein heterodimerization activity and sequence-specific DNA binding transcription factor activity. An important paralog of this gene is TP73.

UniProtKB/Swiss-Prot: P53_HUMAN, P04637
Function: Acts as a tumor suppressor in many tumor types; induces growth arrest or apoptosis depending on the
physiological circumstances and cell type. Involved in cell cycle regulation as a trans-activator that acts to
negatively regulate cell division by controlling a set of genes required for this process. One of the activated
genes is an inhibitor of cyclin-dependent kinases. Apoptosis induction seems to be mediated either by stimulation
of BAX and FAS antigen expression, or by repression of Bcl-2 expression. In cooperation with mitochondrial PPIF
is involved in activating oxidative stress-induced necrosis; the function is largely independent of
transcription. Induces the transcription of long intergenic non-coding RNA p21 (lincRNA-p21) and lincRNA-Mkln1.
LincRNA-p21 participates in TP53-dependent transcriptional repression leading to apoptosis and seem to have to
effect on cell-cycle regulation. Implicated in Notch signaling cross-over. Prevents CDK7 kinase activity when
associated to CAK complex in response to DNA damage, thus stopping cell cycle progression. Isoform 2 enhances the
transactivation activity of isoform 1 from some but not all TP53-inducible promoters. Isoform 4 suppresses
transactivation activity and impairs growth suppression mediated by isoform 1. Isoform 7 inhibits isoform
1-mediated apoptosis

summary for TP53 Gene:
p53 (aka TP53) is a transcription factor whose protein levels and post-translational modification state
alter in response to cellular stress (such as DNA damage, hypoxia, spindle damage). Activation of p53 begins
through a number of mechanisms including phosphorylation by ATM, ATR, Chk1 and MAPKs. MDM2 is a ubiquitn
ligase that binds p53 and targets p53 for proteasomal degradation. Phosphorylation, p14ARF and USP7 prevent
MDM2-p53 interactions, leading to an increase in stable p53 tetramers in the cytoplasm. Further
modifications such as methylation and acetylation lead to an increase in p53 binding to gene specific
response elements. p53 regulates a large number of genes (>100 genes) that control a number of key tumor
suppressing functions such as cell cycle arrest, DNA repair, senescence and apoptosis. Whilst the activation
of p53 often leads to apoptosis, p53 inactivation facilitates tumor progression; inactivating p53 mutations
occur in over 50% of cancers.

Gene Wiki entry for TP53 (P53) Gene


(According to GeneLoc and/or HGNC, and/or
Entrez Gene (NCBI build 37),
and/or miRBase,
Genomic Views according to UCSC (hg19) and Ensembl (release 75), Regulatory elements and Epigenetics data according to QIAGEN, and/or SwitchGear Genomics)
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RefSeq DNA sequence at NCBI GenBank:
NC_000017.10  NC_018928.2  NT_010718.17  
Regulatory elements:
   Regulatory transcription factor binding sites in the TP53 gene promoter:
         CREB   C/EBPbeta   Sp1   ARP-1   
         Other transcription factors

SwitchGear Promoter luciferase reporter plasmids (see all 4): TP53 promoter sequence
   Search Chromatin IP Primers for TP53

Epigenetics:
DNA Methylation CpG Assay Predesigned for Pyrosequencing in human, mouse, rat TP53


Genomic Location:
Genomic View: UCSC Golden Path with GeneCards custom track

Entrez Gene cytogenetic band: 17p13.1   Ensembl cytogenetic band:  17p13.1   HGNC cytogenetic band: 17p13.1

TP53 Gene in genomic location: bands according to Ensembl, locations according to (and/or Entrez Gene and/or Ensembl if different)
TP53 gene location

GeneLoc information about chromosome 17         GeneLoc Exon Structure

GeneLoc location for GC17M007565:  view genomic region     (about GC identifiers)

Start:
7,565,097 bp from pter      End:
7,590,863 bp from pter
Size:
25,767 bases      Orientation:
minus strand
ORGUL member locations:
Legend (see complete legend)

ORGUL Member Locations for TP53

(According to 1UniProtKB, HORDE, 2neXtProt, Ensembl, and/or Reactome, Modification sites according to PhosphoSitePlus, Specific Peptides from DME, RefSeq according to NCBI, PDB rendering according to OCA and/or Proteopedia, Recombinant Proteins from EMD Millipore, R&D Systems, GenScript, Enzo Life Sciences, OriGene, Novus Biologicals, Sino Biological, ProSpec, Cloud-Clone Corp., eBioscience, and/or antibodies-online,
Biochemical Assays by EMD Millipore, R&D Systems, OriGene, GenScript, Cell Signaling Technology, Enzo Life Sciences, Cloud-Clone Corp., eBioscience, and/or antibodies-online, Antibodies by EMD Millipore, R&D Systems, Cell Signaling Technology, OriGene, Novus Biologicals, Thermo Fisher Scientific, Abcam, Cloud-Clone Corp, antibodies-online, and/or others.)
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UniProtKB/Swiss-Prot: P53_HUMAN, P04637 (See protein sequence)
Recommended Name: Cellular tumor antigen p53  
Size: 393 amino acids; 43653 Da
Cofactor: Binds 1 zinc ion per subunit
Subunit: Interacts with AXIN1. Probably part of a complex consisting of TP53, HIPK2 and AXIN1 (By similarity).
Binds DNA as a homotetramer. Interacts with histone acetyltransferases EP300 and methyltransferases HRMT1L2 and
CARM1, and recruits them to promoters. In vitro, the interaction of TP53 with cancer-associated/HPV (E6) viral
proteins leads to ubiquitination and degradation of TP53 giving a possible model for cell growth regulation. This
complex formation requires an additional factor, E6-AP, which stably associates with TP53 in the presence of E6.
Interacts (via C-terminus) with TAF1; when TAF1 is part of the TFIID complex. Interacts with ING4; this
interaction may be indirect. Found in a complex with CABLES1 and TP73. Interacts with HIPK1, HIPK2, and TP53INP1.
Interacts with WWOX. May interact with HCV core protein. Interacts with USP7 and SYVN1. Interacts with HSP90AB1.
Interacts with CHD8; leading to recruit histone H1 and prevent transactivation activity (By similarity).
Interacts with ARMC10, BANP, CDKN2AIP, NUAK1, STK11/LKB1, UHRF2 and E4F1. Interacts with YWHAZ; the interaction
enhances TP53 transcriptional activity. Phosphorylation of YWHAZ on 'Ser-58' inhibits this interaction. Interacts
(via DNA-binding domain) with MAML1 (via N-terminus). Interacts with MKRN1. Interacts with PML (via C-terminus).
Interacts with MDM2; leading to ubiquitination and proteasomal degradation of TP53. Directly interacts with
FBXO42; leading to ubiquitination and degradation of TP53. Interacts (phosphorylated at Ser-15 by ATM) with the
phosphatase PP2A-PPP2R5C holoenzyme; regulates stress-induced TP53-dependent inhibition of cell proliferation.
Interacts with PPP2R2A. Interacts with AURKA, DAXX, BRD7 and TRIM24. Interacts (when monomethylated at Lys-382)
with L3MBTL1. Isoform 1 interacts with isoform 2 and with isoform 4. Interacts with GRK5. Binds to the CAK
complex (CDK7, cyclin H and MAT1) in response to DNA damage. Interacts with CDK5 in neurons. Interacts with
AURKB, SETD2, UHRF2 and NOC2L. Interacts (via N-terminus) with PTK2/FAK1; this promotes ubiquitination by MDM2.
Interacts with PTK2B/PYK2; this promotes ubiquitination by MDM2. Interacts with PRKCG. Interacts with PPIF; the
association implicates preferentially tetrameric TP53, is induced by oxidative stress and is impaired by
cyclosporin A (CsA). Interacts with human cytomegalovirus/HHV-5 protein UL123. Interacts with SNAI1; the
interaction induces SNAI1 degradation via MDM2-mediated ubiquitination and inhibits SNAI1-induced cell invasion.
Interacts with KAT6A. Interacts with UBC9. Interacts with ZNF385B; the interaction is direct. Interacts (via
DNA-binding domain) with ZNF385A; the interaction is direct and enhances p53/TP53 transactivation functions on
cell-cycle arrest target genes, resulting in growth arrest. Interacts with ANKRD2. Interacts with RFFL (via
RING-type zinc finger); involved in p53/TP53 ubiquitination
Selected PDB 3D structures from and Proteopedia for TP53 (see all 119):
1A1U (3D)        1AIE (3D)        1C26 (3D)        1DT7 (3D)        1GZH (3D)        1H26 (3D)    
Secondary accessions: Q15086 Q15087 Q15088 Q16535 Q16807 Q16808 Q16809 Q16810 Q16811 Q16848
Q2XN98 Q3LRW1 Q3LRW2 Q3LRW3 Q3LRW4 Q3LRW5 Q86UG1 Q8J016 Q99659 Q9BTM4 Q9HAQ8 Q9NP68 Q9NPJ2
Q9NZD0 Q9UBI2 Q9UQ61
Alternative promoter usage, Alternative splicing: 9 isoforms:  P04637-1   P04637-2   P04637-3   P04637-4   P04637-5   P04637-6   P04637-7   P04637-8   
P04637-9   (Produced by alternative promoter usage and alternative splicing)

Explore the universe of human proteins at neXtProt for TP53: NX_P04637

Explore proteomics data for TP53 at MOPED

Post-translational modifications: 

  • Acetylated. Acetylation of Lys-382 by CREBBP enhances transcriptional activity. Deacetylation of Lys-382 by SIRT1
    impairs its ability to induce proapoptotic program and modulate cell senescence1
  • Phosphorylation on Ser residues mediates transcriptional activation. Phosphorylated by HIPK1 (By similarity).
    Phosphorylation at Ser-9 by HIPK4 increases repression activity on BIRC5 promoter. Phosphorylated on Thr-18 by
    VRK1. Phosphorylated on Ser-20 by CHEK2 in response to DNA damage, which prevents ubiquitination by MDM2.
    Phosphorylated on Ser-20 by PLK3 in response to reactive oxygen species (ROS), promoting p53/TP53-mediated
    apoptosis. Phosphorylated on Thr-55 by TAF1, which promotes MDM2-mediated degradation. Phosphorylated on Ser-33
    by CDK7 in a CAK complex in response to DNA damage. Phosphorylated on Ser-46 by HIPK2 upon UV irradiation.
    Phosphorylation on Ser-46 is required for acetylation by CREBBP. Phosphorylated on Ser-392 following UV but not
    gamma irradiation. Phosphorylated on Ser-15 upon ultraviolet irradiation; which is enhanced by interaction with
    BANP. Phosphorylated by NUAK1 at Ser-15 and Ser-392; was intially thought to be mediated by STK11/LKB1 but it was
    later shown that it is indirect and that STK11/LKB1-dependent phosphorylation is probably mediated by downstream
    NUAK1 (PubMed:21317932). It is unclear whether AMP directly mediates phosphorylation at Ser-15. Phosphorylated on
    Thr-18 by isoform 1 and isoform 2 of VRK2. Phosphorylation on Thr-18 by isoform 2 of VRK2 results in a reduction
    in ubiquitination by MDM2 and an increase in acetylation by EP300. Stabilized by CDK5-mediated phosphorylation in
    response to genotoxic and oxidative stresses at Ser-15, Ser-33 and Ser-46, leading to accumulation of p53/TP53,
    particularly in the nucleus, thus inducing the transactivation of p53/TP53 target genes. Phosphorylated by DYRK2
    at Ser-46 in response to genotoxic stress. Phosphorylated at Ser-315 and Ser-392 by CDK2 in response to
    DNA-damage1
  • Dephosphorylated by PP2A-PPP2R5C holoenzyme at Thr-55. SV40 small T antigen inhibits the dephosphorylation by the
    AC form of PP2A1
  • May be O-glycosylated in the C-terminal basic region. Studied in EB-1 cell line1
  • Ubiquitinated by MDM2 and SYVN1, which leads to proteasomal degradation. Ubiquitinated by RFWD3, which works in
    cooperation with MDM2 and may catalyze the formation of short polyubiquitin chains on p53/TP53 that are not
    targeted to the proteasome. Ubiquitinated by MKRN1 at Lys-291 and Lys-292, which leads to proteasomal
    degradation. Deubiquitinated by USP10, leading to its stabilization. Ubiquitinated by TRIM24 and RFFL, which
    leads to proteasomal degradation. Ubiquitination by TOPORS induces degradation. Deubiquitination by USP7, leading
    to stabilization. Isoform 4 is monoubiquitinated in an MDM2-independent manner1
  • Monomethylated at Lys-372 by SETD7, leading to stabilization and increased transcriptional activation.
    Monomethylated at Lys-370 by SMYD2, leading to decreased DNA-binding activity and subsequent transcriptional
    regulation activity. Lys-372 monomethylation prevents interaction with SMYD2 and subsequent monomethylation at
    Lys-370. Dimethylated at Lys-373 by EHMT1 and EHMT2. Monomethylated at Lys-382 by SETD8, promoting interaction
    with L3MBTL1 and leading to repress transcriptional activity. Dimethylation at Lys-370 and Lys-382 diminishes p53
    ubiquitination, through stabilizing association with the methyl reader PHF20. Demethylation of dimethylated
    Lys-370 by KDM1A prevents interaction with TP53BP1 and represses TP53-mediated transcriptional activation1
  • Sumoylated with SUMO1. Sumoylated at Lys-386 by UBC91
  • Ubiquitination2 at Lys101, Lys132, Lys164, Lys291, Lys292, Lys305, Lys320, Lys321
  • Modification sites at PhosphoSitePlus

  • See TP53 Protein Expression from SPIRE MOPED, PaxDB, and MaxQB

    REFSEQ proteins (15 alternative transcripts): 
    NP_000537.3  NP_001119584.1  NP_001119585.1  NP_001119586.1  NP_001119587.1  NP_001119588.1  NP_001119589.1  NP_001119590.1  
    NP_001263624.1  NP_001263625.1  NP_001263626.1  NP_001263627.1  NP_001263628.1  NP_001263689.1  NP_001263690.1  

    ENSEMBL proteins: 
     ENSP00000410739   ENSP00000352610   ENSP00000269305   ENSP00000398846   ENSP00000391127  
     ENSP00000391478   ENSP00000458393   ENSP00000425104   ENSP00000423862   ENSP00000424104  
     ENSP00000473895   ENSP00000426252  
    Reactome Protein details: P04637

    TP53 Human Recombinant Protein Products:

    EMD Millipore Purified and/or Recombinant TP53 Protein
    R&D Systems Recombinant & Natural Proteins for TP53 (p53)
    Enzo Life Sciences proteins for TP53
    OriGene Purified Proteins for TP53
    OriGene Protein Over-expression Lysate for TP53
    OriGene MassSpec for TP53
    OriGene Custom Protein Services for TP53
    GenScript Custom Purified and Recombinant Proteins Services for TP53
    Novus Biologicals TP53 Proteins
    Novus Biologicals TP53 Lysates
    Sino Biological Recombinant Protein for TP53
    Browse Sino Biological Cell Lysates
    ProSpec Recombinant Protein for TP53
    Cloud-Clone Corp. Proteins for TP53

     
    Search eBioscience for Proteins for TP53 

     
    antibodies-online proteins for TP53 (61 products) 

     
    antibodies-online peptides for TP53

    TP53 Antibody Products:

    EMD Millipore Mono- and Polyclonal Antibodies for the study of TP53
    R&D Systems Antibodies for TP53 (p53)
    Cell Signaling Technology (CST) Antibodies for TP53  (p53)
    OriGene Antibodies for TP53
    OriGene Custom Antibody Services for TP53
    Novus Biologicals TP53 Antibodies
    Abcam antibodies for TP53
    Cloud-Clone Corp. Antibodies for TP53
    ThermoFisher Antibody for TP53
    antibodies-online antibodies for TP53 (1437 products) 

    TP53 Assay Products:

    EMD Millipore Kits and Assays for the Analysis of TP53
    OriGene Custom Assay Services for TP53
    R&D Systems ELISAs for TP53 (p53)         (see all)
    GenScript Custom Assay Services for TP53
    Cell Signaling Technology (CST) Sandwich ELISA Kits for TP53 (p53)
    Enzo Life Sciences assays for TP53
    Cloud-Clone Corp. ELISAs for TP53
    Cloud-Clone Corp. CLIAs for TP53
    eBioscience ELISAs for TP53: (see all 4
                        p53 (Total/Phospho) InstantOne ELISA 96 tests
    antibodies-online kits for TP53 (80 products) 


    (According to HGNC, IUPHAR, InterPro, ProtoNet, UniProtKB, and/or BLOCKS, Sets of similar genes according to GeneDecks)
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    Selected InterPro protein domains (see all 6):
     IPR011615 p53_DNA-bd
     IPR012346 p53/RUNT-type_TF_DNA-bd
     IPR002117 p53_tumour_suppressor
     IPR008967 p53-like_TF_DNA-bd
     IPR013872 p53_transactivation_domain

    Graphical View of Domain Structure for InterPro Entry P04637

    ProtoNet protein and cluster: P04637

    1 Blocks protein domain: IPB010991 p53

    UniProtKB/Swiss-Prot: P53_HUMAN, P04637
    Domain: The nuclear export signal acts as a transcriptional repression domain. The TADI and TADII motifs (residues
    17 to 25 and 48 to 56) correspond both to 9aaTAD motifs which are transactivation domains present in a large
    number of yeast and animal transcription factors
    Similarity: Belongs to the p53 family


    Find genes that share domains with TP53           About GenesLikeMe


    (According to 1UniProtKB, Genatlas, LifeMap Discovery™, IUBMB, and/or 2DME, Human phenotypes from GenomeRNAi, Animal models from MGI Mar 06 2013, genOway,
    transcription factor targeting from QIAGEN and/or HOMER, miRNA Gene Targets from miRTarBase, shRNA from OriGene, siRNAs from OriGene, QIAGEN, microRNA from QIAGEN, SwitchGear Genomics, Gene Editing from DNA2.0, Clones from OriGene, GenScript, Sino Biological, DNA2.0, Vector BioLabs and/or Addgene, Cell Lines from GenScript, ESI BIO, In Situ Hybridization Assays from Advanced Cell Diagnostics, Flow cytometry from eBioscience, Ontologies according to Gene Ontology Consortium 01 Apr 2014 via Entrez Gene.)
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    Molecular Function:

         UniProtKB/Swiss-Prot Summary: P53_HUMAN, P04637
    Function: Acts as a tumor suppressor in many tumor types; induces growth arrest or apoptosis depending on the
    physiological circumstances and cell type. Involved in cell cycle regulation as a trans-activator that acts to
    negatively regulate cell division by controlling a set of genes required for this process. One of the activated
    genes is an inhibitor of cyclin-dependent kinases. Apoptosis induction seems to be mediated either by stimulation
    of BAX and FAS antigen expression, or by repression of Bcl-2 expression. In cooperation with mitochondrial PPIF
    is involved in activating oxidative stress-induced necrosis; the function is largely independent of
    transcription. Induces the transcription of long intergenic non-coding RNA p21 (lincRNA-p21) and lincRNA-Mkln1.
    LincRNA-p21 participates in TP53-dependent transcriptional repression leading to apoptosis and seem to have to
    effect on cell-cycle regulation. Implicated in Notch signaling cross-over. Prevents CDK7 kinase activity when
    associated to CAK complex in response to DNA damage, thus stopping cell cycle progression. Isoform 2 enhances the
    transactivation activity of isoform 1 from some but not all TP53-inducible promoters. Isoform 4 suppresses
    transactivation activity and impairs growth suppression mediated by isoform 1. Isoform 7 inhibits isoform
    1-mediated apoptosis
    Induction: Up-regulated in response to DNA damage. Isoform 2 is not induced in tumor cells in response to stress

         Genatlas biochemistry entry for TP53:
    tumor suppressor protein p53 required for G1 growth arrest by WAF1 (CDKN1A),following DNA damage or induction of
    apoptosis,also regulating a G2 checkpoint through cyclin B1,transcriptional activator through acetylation of
    transactivation site by CREBBP binding MDM2 resulting in transcriptional silencing and ubiquitin/proteasome
    dependent degradation of p53,activated by conjugation to UBL1 (SUMO1),putative up-regulated c-MYC target
    gene,putative teratologic suppressor gene and modulator of TFIIH (GTF2H),associated in nucleotide excision
    repair,activated by ATM in association with 14.3.3 proteins (YWHA*),tumor suppressor gene (see TSG17A),mutated in
    cancers such as pancreas and endometrial carcinomas,in Barrett's adenocarcinoma (and esophageal squamous cell
    carcinoma),in hepatocellular carcinoma with poor prognosis

         Gene Ontology (GO): Selected molecular function terms (see all 30):    About this table

    GO IDQualified GO termEvidencePubMed IDs
    GO:0000979RNA polymerase II core promoter sequence-specific DNA binding IEA--
    GO:0001077RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in positive regulation of transcription IEA--
    GO:0001085RNA polymerase II transcription factor binding IPI18549481
    GO:0001228RNA polymerase II transcription regulatory region sequence-specific DNA binding transcription factor activity involved in positive regulation of transcription IDA17146433
    GO:0002020protease binding IPI11923872
         
    Find genes that share ontologies with TP53           About GenesLikeMe


    Phenotypes:
         4 GenomeRNAi human phenotypes for TP53:
     Decreased viability of wild-ty  Enable proliferation in B-Raf   Increased cell death HMECs cel  Increased resistance to MDM2 i 

         Selected MGI mutant phenotypes (inferred from 73 alleles(MGI details for Trp53) (see all 27):
     adipose tissue  behavior/neurological  cardiovascular system  cellular  craniofacial 
     digestive/alimentary  embryogenesis  endocrine/exocrine gland  growth/size/body  hematopoietic system 
     homeostasis/metabolism  immune system  integument  limbs/digits/tail  liver/biliary system 

    Find genes that share phenotypes with TP53           About GenesLikeMe

    Animal Models:
         MGI mouse knock-outs for TP53: Trp53tm1Mlh Trp53tm3Wahl Trp53tm5Wahl Trp53tm2Wahl Trp53tm1.1Manf Trp53tm1Tyj
                                                         Trp53tm1Mok Trp53tm1Sia Trp53tm1.1Brn Trp53tm1Brd

       genOway: Develop your customized and physiologically relevant rodent model for TP53

    Transcription Factor Targeting: 
    1892 GeneGlobe predicted Target genes for TP53 (see first 10):
    AATF,  ABCC3,  ABCC6P2,  ABCD1,  ABCF1,  ABHD12,  ABHD15,  ABHD4,  ABHD8,  ABTB2,
    ACP1,  ACRBP,  ACSM3,  ACTB,  ACTG1,  ACTN3,  ACTR2,  ACTR5,  ADAMTS9,  ADARB1,
    ADCYAP1R1,  ADH1B,  ADPRHL1,  ADRB2,  ADSSL1,  AGA,  AGAP3,  AGAP7,  AGBL1,  AGPAT9,
    AGRN,  AIF1L,  AIPL1,  AK1,  AKAP8,  AKR1C1,  AKR1C3,  AKTIP,  ALDH2,  ALDH5A1,
    ALG1L,  ALK,  ALMS1P,  ALOX12P2,  ALOX5,  AMDHD2,  AMH,  AMPH,  ANK2,  ANKDD1A,
    ANKFY1,  ANKK1,  ANKRD13B,  ANKRD13D,  ANKRD23,  ANKRD33B,  ANTXR1,  ANTXR2,  ANXA13,  ANXA2P3,
    AOC2,  AOC3,  AP1M1,  AP1M2,  APBA2,  APBB3,  APC2,  APH1B,  APLP1,  APLP2,
    APOF,  AQP2,  AQP5,  AQP6,  ARG1,  ARHGAP12,  ARHGAP29,  ARHGAP30,  ARHGEF2,  ARHGEF3,
    ARHGEF5,  ARHGEF7,  ARID1A,  ARID3A,  ARID3C,  ARL1,  ARL2,  ARL5B,  ARL6IP5,  ARSG,
    ARSH,  ART3,  ASAH2,  ASMT,  ASPSCR1,  ASTL,  ASXL2,  ATAD2,  ATG2A,  ATG5,
    ATOH1,  ATOX1,  ATP13A2,  ATP1A1,  ATP2A1,  ATP5E,  ATP5SL,  ATP6V0C,  ATP7B,  ATP8B3,
    ATRIP,  B3GALT4,  B3GALT5,  B3GNT9,  B4GALNT1,  BACE1,  BACE2,  BAG1,  BAIAP2L1,  BAK1,
    BANP,  BARD1,  BCAM,  BCAN,  BCL2L12,  BCL7C,  BCR,  BECN1,  BEND3,  BGLAP,
    BHLHE40,  BIRC2,  BLM,  BLNK,  BLZF1,  BMP1,  BMP6,  BMP8A,  BNC1,  BNIP3,
    BOC,  BOLA3,  BRAF,  BRD4,  BTG2,  BTG3,  BTLA,  BTN1A1,  BTN2A1,  BTN2A2,
    BTN3A1,  BTN3A3,  BUB3,  C10orf10,  C10orf2,  C10orf99,  C11orf1,  C11orf85,  C12orf49,  C12orf76,
    C16orf78,  C16orf87,  C16orf89,  C17orf72,  C17orf82,  C17orf89,  C17orf99,  C18orf56,  C19orf24,  C19orf35,
    C19orf38,  C19orf67,  C1orf147,  C1orf204,  C1orf74,  C21orf33,  C2CD2L,  C2orf66,  C2orf70,  C3orf27,
    C3orf62,  C4B,  C4orf36,  C5orf24,  C6orf106,  C7orf49,  C8G,  C9orf153,  C9orf78,  CA5A,
    CA5B,  CA7,  CABLES2,  CACNA1E,  CADM1,  CALML5,  CALR,  CAPN2,  CARTPT,  CASKIN1,
    CASP10,  CASP12,  CASP14,  CAST,  CBLN4,  CBX3,  CC2D1B,  CCDC11,  CCDC120,  CCDC127,
    CCDC148,  CCDC157,  CCDC42,  CCDC51,  CCDC59,  CCDC61,  CCDC85B,  CCL18,  CCL19,  CCL21,
    CCL27,  CCL28,  CCNB1,  CCNK,  CCNYL2,  CCR5,  CCT6A,  CCT6B,  CD200,  CD27,
    CD2BP2,  CD38,  CD3E,  CD3EAP,  CD68,  CD7,  CD79A,  CD8A,  CDAN1,  CDC123,
    CDC20,  CDC37L1,  CDC42EP2,  CDCA5,  CDH1,  CDH13,  CDH19,  CDH22,  CDIPT,  CDK5,
    CDK5R1,  CDK5RAP3,  CDK9,  CDKN2B,  CDKN2D,  CDT1,  CEACAM6,  CEBPZ,  CELSR3,  CEND1,
    CEP164,  CEP78,  CETN1,  CETP,  CFI,  CFL1,  CFLAR,  CGB7,  CGB8,  CHAC1,
    CHAD,  CHCHD10,  CHD9,  CHEK1,  CHID1,  CHMP6,  CHRFAM7A,  CHRNA10,  CHRNA4,  CHST2,
    CHTF18,  CIDECP,  CIRBP,  CISD1,  CKMT2,  CKS1B,  CLC,  CLCN6,  CLDN17,  CLEC3A,
    CLECL1,  CLIC1,  CLIP1,  CLK2,  CLSTN2,  CMTM5,  CNN3,  CNNM2,  CNP,  COG7,
    COG8,  COL18A1,  COL8A1,  COMMD6,  COPB1,  COPB2,  COPS3,  COPS8,  COQ7,  CORO2A,
    CORT,  COX5A,  COX5B,  COX6A1,  CPN1,  CPO,  CPSF3,  CPT1C,  CRCT1,  CREBBP,
    CREG1,  CRNKL1,  CRYGS,  CRYZL1,  CSAG2,  CSK,  CSN2,  CTCF,  CTF1,  CTNNA3,
    CTNNB1,  CTR9,  CTRL,  CTSW,  CUEDC2,  CUL4B,  CXCL9,  CXXC1,  CYB5R2,  CYBA,
    CYLD,  CYMP,  CYP1A1,  CYP1A2,  CYP21A2,  CYP27A1,  CYP2C19,  CYP4F2,  CYP4F22,  CYTL1,
    DAB2IP,  DACT3,  DAD1,  DAK,  DAP,  DAPK3,  DAPL1,  DAXX,  DAZ1,  DAZ2,
    DAZ3,  DAZ4,  DAZAP2,  DCBLD2,  DCC,  DCK,  DCLRE1A,  DCLRE1C,  DCP1A,  DCTN3,
    DDA1,  DDAH2,  DDIT4,  DDR2,  DDT,  DDX3X,  DDX5,  DDX50,  DDX52,  DECR1,
    DEDD,  DEFB1,  DEFB126,  DEFB127,  DES,  DFNA5,  DGCR5,  DGKA,  DGKQ,  DHCR24,
    DHFR,  DHPS,  DHX30,  DHX33,  DHX38,  DIABLO,  DIRC2,  DIS3,  DKFZP586I1420,  DLEC1,
    DLL4,  DMBT1,  DMP1,  DMRTB1,  DMTF1,  DNAH3,  DNAH8,  DNAJB1,  DNAJB2,  DNAJB3,
    DNAJC5,  DNAJC7,  DNALI1,  DNASE1L2,  DND1,  DNMT3A,  DPAGT1,  DPEP3,  DPY19L2P2,  DRAP1,
    DRG1,  DSTYK,  DTL,  DTX1,  DUOX1,  DUOX2,  DUOXA1,  DUSP2,  DUSP3,  DUSP4,
    DUSP5,  DUT,  DYRK2,  DZIP1,  E2F3,  E2F4,  E4F1,  EAF2,  EARS2,  EBP,
    ECD,  EDN1,  EED,  EEF1G,  EEF2,  EFEMP2,  EFNA2,  EFNA3,  EFS,  EGFL6,
    EGFL8,  EGFR,  EHD1,  EHMT1,  EI24,  EID1,  EID2,  EID2B,  EID3,  EIF2AK4,
    EIF2B3,  EIF2S1,  EIF4A1,  EIF4A3,  EIF4E1B,  EIF4G1,  EIF5A2,  ELF1,  ELFN2,  ELK4,
    ELMO3,  ELOF1,  ELOVL1,  ENO2,  ENSA,  EOMES,  EPB41L3,  EPC1,  EPDR1,  EPHA10,
    EPHA5,  EPHB4,  EPHX1,  EPN3,  EPOR,  ERAL1,  ERBB4,  ERCC2,  ERCC3,  ERCC4,
    ERCC5,  EREG,  ERF,  ESP33,  ETAA1,  ETHE1,  ETS2,  EVI5,  EXTL3,  FAF1,
    FAHD1,  FAM109A,  FAM126A,  FAM153C,  FAM157C,  FAM166A,  FAM171A2,  FAM179A,  FAM180A,  FAM19A3,
    FAM3A,  FAM41C,  FAM57A,  FAM64A,  FAM73B,  FAM83D,  FAM83E,  FAM83H,  FAM89B,  FAM8A1,
    FAM90A1,  FAM92A1,  FANCC,  FANCD2,  FARSA,  FASTK,  FBF1,  FBXL15,  FBXL18,  FBXL8,
    FBXO11,  FBXO5,  FBXW5,  FBXW8,  FBXW9,  FCGR1A,  FCGR2A,  FCN2,  FEM1B,  FER,
    FERD3L,  FGF17,  FGGY,  FGR,  FH,  FHOD3,  FIBP,  FICD,  FIGF,  FIZ1,
    FLCN,  FLII,  FLJ26850,  FLJ30403,  FLJ37453,  FLJ42627,  FLJ46010,  FMNL2,  FMO2,  FMO6P,
    FNBP1L,  FOSL1,  FOSL2,  FOXD4L1,  FOXD4L2,  FOXD4L3,  FOXE1,  FOXF1,  FOXH1,  FOXO3B,
    FOXR1,  FPGS,  FPGT,  FREM2,  FSCN1,  FSCN2,  FTSJ1,  FUK,  FUT11,  FUT2,
    FXYD3,  G3BP1,  G6PD,  GABRA2,  GABRA3,  GABRQ,  GADD45G,  GADD45GIP1,  GAL,  GALM,
    GALNT10,  GALNT6,  GALNT9,  GALT,  GAPDH,  GAR1,  GAS6,  GAS7,  GCA,  GCHFR,
    GCM2,  GCNT2,  GCNT3,  GCOM1,  GDA,  GFER,  GGNBP1,  GIMAP1,  GIMAP5,  GJB1,
    GLA,  GLI1,  GLIPR1,  GLIPR2,  GLO1,  GLTSCR2,  GLUD2,  GLYAT,  GMIP,  GML,
    GMNN,  GMPS,  GNA11,  GNAI1,  GNAT3,  GNB2L1,  GNL1,  GNPTAB,  GOT1L1,  GPLD1,
    GPNMB,  GPR111,  GPR12,  GPR132,  GPR133,  GPR173,  GPR176,  GPR18,  GPR84,  GPR88,
    GPRC5A,  GPRIN2,  GPSM2,  GPT,  GPX1,  GPX2,  GPX4,  GRB2,  GREM1,  GRIA2,
    GRIA3,  GRID2IP,  GRIN2D,  GSK3A,  GSPT1,  GSR,  GSTO2,  GSTP1,  GSTT2,  GSTTP2,
    GTF2A1L,  GTF2H4,  GUK1,  GZMH,  H19,  H2AFX,  HABP4,  HAGH,  HAPLN2,  HARS2,
    HAS1,  HCG18,  HCK,  HCN3,  HDAC8,  HEG1,  HELQ,  HERC2,  HERC3,  HES4,
    HEXA,  HEXDC,  HEY1,  HIATL2,  HLA-DRB1,  HLA-J,  HMOX1,  HMX2,  HMX3,  HN1,
    HNF1A,  HNRNPA2B1,  HNRNPA3P1,  HNRNPD,  HNRNPH2,  HNRNPUL1,  HOOK2,  HOXA10,  HOXA11,  HOXA13,
    HOXA6,  HOXA7,  HOXA9,  HOXB13,  HPRT1,  HR,  HRG,  HRK,  HS3ST6,  HSBP1,
    HSD17B10,  HSD17B13,  HSD17B4,  HSF4,  HSP90AA1,  HSPA14,  HSPA2,  HSPA6,  HSPA7,  HSPBAP1,
    HSPE1,  HTR3C,  HYOU1,  ID2,  IDUA,  IFNGR1,  IGF2,  IGFBP1,  IKBKB,  IKBKG,
    IL11RA,  IL17C,  IL17RD,  IL1RAPL1,  IL1RL1,  IL25,  IL2RA,  IL7,  IL9R,  IMP3,
    INE1,  INGX,  INHA,  INHBE,  INO80B,  INSR,  INSRR,  INTS6,  INTS7,  INTS8,
    IP6K3,  IPMK,  IQCB1,  IQCE,  IQCF3,  IREB2,  IRF3,  IRF4,  IRF5,  IRF6,
    IRX4,  ISG15,  ITCH,  ITGA6,  ITGAD,  ITGB1BP1,  ITPKB,  ITSN1,  IWS1,  JAG1,
    JPH1,  JSRP1,  JUP,  KANK2,  KCNA6,  KCNA7,  KCNK7,  KCNN4,  KCNQ4,  KCNT1,
    KCP,  KCTD21,  KDELR1,  KIAA0895L,  KIAA0947,  KIAA1279,  KIAA1755,  KIAA1919,  KIF17,  KIF18A,
    KIF22,  KIFC3,  KIR2DS4,  KIR3DL2,  KIRREL2,  KISS1R,  KIT,  KL,  KLF13,  KLF5,
    KLF6,  KLHDC9,  KLHL14,  KLK14,  KMO,  KPNA3,  KPNA5,  KRI1,  KRT1,  KRT13,
    KRT15,  KRT17,  KRT3,  KRT35,  KRT36,  KRT5,  KRT7,  KRT74,  KRT76,  KRT77,
    KRT84,  LAMC3,  LAMP3,  LATS1,  LATS2,  LCE1B,  LCE1C,  LCE1D,  LCE5A,  LCN10,
    LCN12,  LCN6,  LDHD,  LETM1,  LETMD1,  LEUTX,  LGALS1,  LGALS7,  LGALS7B,  LGI3,
    LHX1,  LIG3,  LIN37,  LINGO4,  LIPE,  LMBR1L,  LMCD1,  LNP1,  LOC100128035,  LOC100128430,
    LOC100129055,  LOC100129060,  LOC100129637,  LOC100130100,  LOC100130238,  LOC100130938,  LOC100131257,  LOC100131582,  LOC100132272,  LOC100286925,
    LOC100286986,  LOC100287294,  LOC100287329,  LOC100287525,  LOC100287951,  LOC100288123,  LOC100288181,  LOC100288482,  LOC100288656,  LOC100288802,
    LOC100288869,  LOC100288893,  LOC100289283,  LOC113230,  LOC128322,  LOC143666,  LOC220077,  LOC284889,  LOC342293,  LOC344887,
    LOC388955,  LOC391578,  LOC392196,  LOC400499,  LOC407835,  LOC439951,  LOC440031,  LOC440300,  LOC553137,  LOC642423,
    LOC642515,  LOC642635,  LOC642648,  LOC643043,  LOC644172,  LOC646938,  LOC646999,  LOC653486,  LOC723805,  LOC728989,
    LOC729275,  LOC729324,  LOC729654,  LOC729681,  LOC729856,  LOC729941,  LOC730974,  LONRF2,  LOXHD1,  LPAR2,
    LPO,  LPP,  LRRC14,  LRRC26,  LRRC3,  LRRC4,  LRRC48,  LRRC66,  LRRC8B,  LRWD1,
    LSM11,  LSR,  LSS,  LST1,  LTA,  LTB,  LTBP2,  LY6G6C,  LYPD1,  LYPD2,
    LYPD4,  LYPD5,  LYST,  LYZ,  LZTS1,  MAGEA12,  MAGEA2,  MAGEA2B,  MAGEA3,  MAGEA4,
    MAGEA6,  MAGEA8,  MAGEC1,  MAGEC3,  MAGED2,  MAGEH1,  MANBA,  MAP3K1,  MAP3K5,  MAP3K6,
    MAP4K2,  MAPK11,  MAPK12,  MAPK8IP1,  MAPKAPK5,  MARVELD3,  MAS1,  MATN2,  MAZ,  MBD1,
    MBD2,  MBL2,  MCAM,  MCCC2,  MCHR1,  MCM2,  MCM4,  MCM5,  MCM8,  MCTS1,
    MDC1,  MDGA2,  ME1,  MED20,  MED24,  MED28,  MED29,  MEIG1,  MET,  MEX3A,
    MEX3D,  MFAP2,  MFSD3,  MGC34034,  MGC57346,  MIB1,  MIF,  MKS1,  MLANA,  MMP11,
    MMP23B,  MMP26,  MMP7,  MNT,  MOGAT3,  MON2,  MORF4L1,  MORN1,  MPDU1,  MPP2,
    MPV17,  MRGPRX2,  MRPL13,  MRPL18,  MRPL34,  MRPL46,  MRPS11,  MRPS18C,  MRPS24,  MRTO4,
    MSH3,  MSH5,  MSI1,  MSLN,  MSN,  MSRA,  MSX1,  MSX2P1,  MT1A,  MT1DP,
    MT1M,  MT2A,  MT3,  MTA1,  MTA2,  MTBP,  MTERFD3,  MTF1,  MTHFD2,  MTHFD2L,
    MTHFR,  MUC2,  MUC5AC,  MUC5B,  MUC6,  MUM1,  MVD,  MVP,  MYADML,  MYADML2,
    MYBL2,  MYCN,  MYCNOS,  MYL3,  MYL9,  MYLK,  MYLPF,  MYNN,  MYO1H,  MYO6,
    MYO7B,  MYOG,  NACA2,  NAE1,  NALCN,  NANOG,  NARFL,  NAT2,  NAT8,  NBL1,
    NBN,  NBPF11,  NBPF3,  NBR1,  NCF1,  NCOA6,  NDEL1,  NDNL2,  NDST1,  NDUFAF1,
    NDUFB11,  NDUFS8,  NDUFV2,  NEFH,  NFKB2,  NFX1,  NFXL1,  NFYA,  NHLRC1,  NHLRC2,
    NHS,  NHSL1,  NINJ1,  NIP7,  NIT1,  NKAP,  NKAPL,  NKD2,  NKIRAS2,  NKX2-4,
    NKX3-1,  NLE1,  NMBR,  NMD3,  NME7,  NOD1,  NOL12,  NOL3,  NOMO1,  NOMO2,
    NOSIP,  NOTCH4,  NOTUM,  NOVA2,  NOX4,  NPBWR2,  NPFF,  NPFFR1,  NPHS1,  NPY1R,
    NPY6R,  NQO1,  NR2C1,  NRXN3,  NSUN6,  NT5C,  NT5M,  NTF4,  NTHL1,  NTN1,
    NTN5,  NTNG1,  NTRK1,  NUCKS1,  NUDT16,  NUDT5,  NUMBL,  NUP93,  NXF4,  NXT1,
    OAZ1,  OBSL1,  OCA2,  OCM,  ODF3,  ODF3L2,  OGDH,  OPA1,  OPN1MW,  OPN1SW,
    OR10K1,  OR13C9,  OR1E1,  OR1N2,  OR2A9P,  OR2M2,  OR4C46,  OR4D1,  OR4D2,  OR51A2,
    OR51I1,  OR51I2,  OR51Q1,  OR52B6,  OR52H1,  OR6F1,  ORAI1,  OTUB2,  OXER1,  P2RX2,
    P2RX6,  PABPC3,  PACS2,  PAICS,  PAK1,  PANX3,  PAQR6,  PARD6G,  PARVG,  PBX4,
    PBXIP1,  PC,  PCDH11X,  PCDH11Y,  PCDHB1,  PDC,  PDCD1LG2,  PDCD5,  PDCL3,  PDE6A,
    PDE8A,  PDF,  PDK1,  PDS5B,  PDXP,  PEAR1,  PECR,  PEMT,  PER1,  PERP,
    PES1,  PEX14,  PFDN2,  PFDN6,  PFN2,  PFN4,  PGBD1,  PGC,  PGF,  PGP,
    PHC2,  PHEX,  PHF21B,  PHLDB2,  PI3,  PIBF1,  PIF1,  PIGV,  PIK3C3,  PIM3,
    PIP,  PISD,  PKD1P1,  PKP4,  PLA2G10,  PLCB2,  PLCD1,  PLD4,  PLEKHA7,  PLEKHG1,
    PLEKHJ1,  PLEKHO2,  PLOD3,  PLSCR5,  PLXNA2,  PLXNB1,  PMP22,  PMPCA,  PMPCB,  PNKP,
    PNLDC1,  PNLIPRP2,  PNO1,  PNPLA2,  PODXL,  POLK,  POLR2A,  POLR2E,  POLR2G,  POLR3K,
    POMT1,  POMZP3,  PON1,  PORCN,  POU1F1,  POU4F2,  PPAPDC2,  PPAT,  PPFIBP1,  PPM1A,
    PPM1F,  PPOX,  PPP1CA,  PPP1R12C,  PPP1R13B,  PPP1R13L,  PPP1R16A,  PPP2R5A,  PPP2R5B,  PPP3CB,
    PPP4R1,  PPY,  PRAMEF10,  PRAMEF13,  PRAMEF22,  PRAMEF25,  PRAMEF4,  PRAMEF6,  PRAMEF8,  PRDM1,
    PRDM7,  PRICKLE4,  PRINS,  PRKCD,  PRKDC,  PRMT1,  PRNP,  PROM1,  PROM2,  PROP1,
    PRPS1,  PRR12,  PRR15,  PRRG2,  PRRT1,  PRRT2,  PRX,  PSD2,  PSEN2,  PSENEN,
    PSMA1,  PSMB10,  PSMC4,  PSMD11,  PSMD2,  PSMD5,  PSME3,  PSME4,  PSMG3,  PSPH,
    PSRC1,  PTCD3,  PTCRA,  PTH1R,  PTMA,  PTMS,  PTPN9,  PTPRM,  PUS10,  PVR,
    PVRL4,  PVT1,  PYCARD,  PYGO2,  QPRT,  QSOX2,  R3HDM2,  RAB11B,  RAB27B,  RAB33B,
    RAB35,  RAB39B,  RAB40AL,  RAB6B,  RAD23A,  RAD23B,  RAD50,  RAD51,  RAD52,  RAD54L2,
    RAD9A,  RALA,  RALBP1,  RANBP2,  RANGAP1,  RAPGEF5,  RAPGEFL1,  RARA,  RARRES3,  RASD1,
    RASGRF2,  RASGRP4,  RASSF5,  RBBP4,  RBL1,  RBM10,  RBM17,  RBM19,  RBP1,  RBX1,
    REEP1,  RER1,  RETNLB,  RFT1,  RGMA,  RGS12,  RHBDL1,  RHBDL3,  RHEB,  RHOC,
    RHOF,  RHOH,  RHOT2,  RIBC1,  RIF1,  RIMBP3C,  RIMS1,  RIPK1,  </