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    SIRT2 sirtuin 2 [ Homo sapiens (human) ]

    Gene ID: 22933, updated on 10-Dec-2024

    Summary

    Official Symbol
    SIRT2provided by HGNC
    Official Full Name
    sirtuin 2provided by HGNC
    Primary source
    HGNC:HGNC:10886
    See related
    Ensembl:ENSG00000068903 MIM:604480; AllianceGenome:HGNC:10886
    Gene type
    protein coding
    RefSeq status
    REVIEWED
    Organism
    Homo sapiens
    Lineage
    Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia; Eutheria; Euarchontoglires; Primates; Haplorrhini; Catarrhini; Hominidae; Homo
    Also known as
    SIR2; SIR2L; SIR2L2
    Summary
    This gene encodes a member of the sirtuin family of proteins, homologs to the yeast Sir2 protein. Members of the sirtuin family are characterized by a sirtuin core domain and grouped into four classes. The functions of human sirtuins have not yet been determined; however, yeast sirtuin proteins are known to regulate epigenetic gene silencing and suppress recombination of rDNA. Studies suggest that the human sirtuins may function as intracellular regulatory proteins with mono-ADP-ribosyltransferase activity. The protein encoded by this gene is included in class I of the sirtuin family. Several transcript variants are resulted from alternative splicing of this gene. [provided by RefSeq, Jul 2010]
    Expression
    Ubiquitous expression in brain (RPKM 37.1), ovary (RPKM 15.7) and 25 other tissues See more
    Orthologs
    NEW
    Try the new Gene table
    Try the new Transcript table

    Genomic context

    See SIRT2 in Genome Data Viewer
    Location:
    19q13.2
    Exon count:
    17
    Annotation release Status Assembly Chr Location
    RS_2024_08 current GRCh38.p14 (GCF_000001405.40) 19 NC_000019.10 (38878555..38899618, complement)
    RS_2024_08 current T2T-CHM13v2.0 (GCF_009914755.1) 19 NC_060943.1 (41682611..41703673, complement)
    RS_2024_09 previous assembly GRCh37.p13 (GCF_000001405.25) 19 NC_000019.9 (39369195..39390258, complement)

    Chromosome 19 - NC_000019.10Genomic Context describing neighboring genes Neighboring gene uncharacterized LOC124904712 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 14595 Neighboring gene H3K4me1 hESC enhancer GRCh37_chr19:39333551-39334051 Neighboring gene heterogeneous nuclear ribonucleoprotein L Neighboring gene H3K27ac hESC enhancer GRCh37_chr19:39338887-39339825 Neighboring gene H3K27ac hESC enhancer GRCh37_chr19:39339826-39340763 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 14597 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 14598 Neighboring gene ATAC-STARR-seq lymphoblastoid silent region 10586 Neighboring gene H3K27ac hESC enhancer GRCh37_chr19:39343185-39343829 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 14599 Neighboring gene ATAC-STARR-seq lymphoblastoid silent region 10587 Neighboring gene ReSE screen-validated silencer GRCh37_chr19:39350790-39350958 Neighboring gene Sharpr-MPRA regulatory region 3140 Neighboring gene NANOG-H3K27ac-H3K4me1 hESC enhancer GRCh37_chr19:39356343-39356918 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 14600 Neighboring gene H3K4me1 hESC enhancer GRCh37_chr19:39360061-39360631 Neighboring gene H3K4me1 hESC enhancer GRCh37_chr19:39360661-39361267 Neighboring gene H3K4me1 hESC enhancer GRCh37_chr19:39361268-39361873 Neighboring gene Ras and Rab interactor like Neighboring gene ATAC-STARR-seq lymphoblastoid silent region 10589 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 14602 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 14603 Neighboring gene H3K4me1 hESC enhancer GRCh37_chr19:39384205-39384706 Neighboring gene NFKB inhibitor beta Neighboring gene H3K4me1 hESC enhancer GRCh37_chr19:39402426-39403052 Neighboring gene coiled-coil glutamate rich protein 2

    Genomic regions, transcripts, and products

    Expression

    • Project title: HPA RNA-seq normal tissues
    • Description: RNA-seq was performed of tissue samples from 95 human individuals representing 27 different tissues in order to determine tissue-specificity of all protein-coding genes
    • BioProject: PRJEB4337
    • Publication: PMID 24309898
    • Analysis date: Wed Apr 4 07:08:55 2018

    Bibliography

    GeneRIFs: Gene References Into Functions

    What's a GeneRIF?

    Pathways from PubChem

    Interactions

    Products Interactant Other Gene Complex Source Pubs Description

    General gene information

    Markers

    Clone Names

    • FLJ35621, FLJ37491

    Gene Ontology Provided by GOA

    Function Evidence Code Pubs
    enables DNA-binding transcription factor binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    enables NAD+ binding IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    enables NAD+ binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    NOT enables NAD+-protein ADP-ribosyltransferase activity TAS
    Traceable Author Statement
    more info
    PubMed 
    enables NAD+-protein poly-ADP-ribosyltransferase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    NOT enables NAD+-protein poly-ADP-ribosyltransferase activity TAS
    Traceable Author Statement
    more info
    PubMed 
    enables NAD-dependent protein demyristoylase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables NAD-dependent protein depalmitoylase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables NAD-dependent protein lysine deacetylase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables NAD-dependent protein lysine deacetylase activity IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    enables NAD-dependent protein lysine deacetylase activity TAS
    Traceable Author Statement
    more info
     
    enables chromatin binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables histone H3K14 deacetylase activity, NAD-dependent IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3K18 deacetylase activity, NAD-dependent IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3K4 deacetylase activity, NAD-dependent IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3K56 deacetylase activity, NAD-dependent IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3K9 deacetylase activity, NAD-dependent IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H4K16 deacetylase activity, NAD-dependent IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables histone acetyltransferase binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    enables histone deacetylase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables histone deacetylase activity, NAD-dependent IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    NOT enables histone deacetylase activity, NAD-dependent IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables histone deacetylase activity, NAD-dependent IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables histone deacetylase binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    enables protein binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    enables protein lysine deacetylase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables protein lysine deacetylase activity IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    NOT enables tubulin deacetylase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables tubulin deacetylase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables ubiquitin binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables zinc ion binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    Process Evidence Code Pubs
    involved_in NLRP3 inflammasome complex assembly IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in autophagy IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in cell division IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in cellular response to caloric restriction ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in cellular response to epinephrine stimulus ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in cellular response to hypoxia IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in cellular response to oxidative stress ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in epigenetic regulation of gene expression IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in heterochromatin formation NAS
    Non-traceable Author Statement
    more info
    PubMed 
    involved_in innate immune response IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in lipid catabolic process ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in meiotic cell cycle IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in mitotic nuclear membrane reassembly TAS
    Traceable Author Statement
    more info
     
    involved_in myelination in peripheral nervous system ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in negative regulation of DNA-templated transcription IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in negative regulation of NLRP3 inflammasome complex assembly IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in negative regulation of autophagy IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in negative regulation of fat cell differentiation ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in negative regulation of oligodendrocyte progenitor proliferation ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in negative regulation of peptidyl-threonine phosphorylation ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in negative regulation of protein catabolic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in negative regulation of reactive oxygen species metabolic process ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in negative regulation of satellite cell differentiation ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in negative regulation of striated muscle tissue development IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in negative regulation of transcription by RNA polymerase II IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in negative regulation of transcription by RNA polymerase II IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in peptidyl-lysine deacetylation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of DNA binding ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of attachment of spindle microtubules to kinetochore ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of cell division ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of execution phase of apoptosis ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of fatty acid biosynthetic process IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of meiotic nuclear division ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of oocyte maturation ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of proteasomal ubiquitin-dependent protein catabolic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in positive regulation of proteasomal ubiquitin-dependent protein catabolic process ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of transcription by RNA polymerase II ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in post-translational protein modification IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in proteasome-mediated ubiquitin-dependent protein catabolic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    NOT involved_in protein deacetylation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in protein deacetylation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in rDNA heterochromatin formation IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    involved_in rDNA heterochromatin formation NAS
    Non-traceable Author Statement
    more info
    PubMed 
    involved_in regulation of cell cycle IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in regulation of exit from mitosis NAS
    Non-traceable Author Statement
    more info
    PubMed 
    involved_in regulation of myelination ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in regulation of phosphorylation NAS
    Non-traceable Author Statement
    more info
    PubMed 
    involved_in response to redox state NAS
    Non-traceable Author Statement
    more info
    PubMed 
    involved_in substantia nigra development HEP PubMed 
    involved_in subtelomeric heterochromatin formation NAS
    Non-traceable Author Statement
    more info
    PubMed 
    NOT involved_in tubulin deacetylation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in tubulin deacetylation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in tubulin deacetylation ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    Component Evidence Code Pubs
    located_in Schmidt-Lanterman incisure ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in centriole IDA
    Inferred from Direct Assay
    more info
    PubMed 
    located_in centrosome IDA
    Inferred from Direct Assay
    more info
    PubMed 
    part_of chromatin silencing complex NAS
    Non-traceable Author Statement
    more info
    PubMed 
    located_in chromosome IDA
    Inferred from Direct Assay
    more info
    PubMed 
    located_in chromosome, telomeric region IEA
    Inferred from Electronic Annotation
    more info
     
    located_in cytoplasm IDA
    Inferred from Direct Assay
    more info
    PubMed 
    located_in cytosol IDA
    Inferred from Direct Assay
    more info
    PubMed 
    located_in cytosol TAS
    Traceable Author Statement
    more info
     
    located_in glial cell projection ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in growth cone IEA
    Inferred from Electronic Annotation
    more info
     
    located_in heterochromatin ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in juxtaparanode region of axon ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in lateral loop ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in meiotic spindle ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in microtubule IDA
    Inferred from Direct Assay
    more info
    PubMed 
    located_in midbody IDA
    Inferred from Direct Assay
    more info
    PubMed 
    located_in mitochondrion HTP PubMed 
    located_in mitotic spindle IDA
    Inferred from Direct Assay
    more info
    PubMed 
    located_in myelin sheath ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in nucleolus IDA
    Inferred from Direct Assay
    more info
     
    is_active_in nucleus IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    located_in nucleus IDA
    Inferred from Direct Assay
    more info
    PubMed 
    located_in paranodal junction ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in paranode region of axon ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in perikaryon ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in perinuclear region of cytoplasm ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in plasma membrane IDA
    Inferred from Direct Assay
    more info
     
    located_in spindle IDA
    Inferred from Direct Assay
    more info
    PubMed 

    General protein information

    Preferred Names
    NAD-dependent protein deacetylase sirtuin-2
    Names
    NAD-dependent deacetylase sirtuin-2
    NAD-dependent protein defatty-acylase sirtuin-2
    SIR2-like protein 2
    regulatory protein SIR2 homolog 2
    silent information regulator 2
    sir2-related protein type 2
    sirtuin type 2
    NP_001180215.1
    NP_036369.2
    NP_085096.1
    XP_011524957.1
    XP_047294424.1
    XP_047294425.1
    XP_054176249.1
    XP_054176250.1
    XP_054176251.1
    XP_054187933.1
    XP_054187934.1
    XP_054187935.1

    NCBI Reference Sequences (RefSeq)

    NEW Try the new Transcript table

    RefSeqs maintained independently of Annotated Genomes

    These reference sequences exist independently of genome builds. Explain

    These reference sequences are curated independently of the genome annotation cycle, so their versions may not match the RefSeq versions in the current genome build. Identify version mismatches by comparing the version of the RefSeq in this section to the one reported in Genomic regions, transcripts, and products above.

    mRNA and Protein(s)

    1. NM_001193286.2NP_001180215.1  NAD-dependent protein deacetylase sirtuin-2 isoform 3

      See identical proteins and their annotated locations for NP_001180215.1

      Status: REVIEWED

      Description
      Transcript Variant: This variant (3) lacks an exon in the 5' region, resulting in a downstream in-frame AUG start codon, and also lacks a big segment in the 3' region, resulting in a frame-shift, as compared to variant 1. The resulting isoform (3) has a shorter N-terminus, and a shorter and different C-terminus, as compared to isoform 1.
      Source sequence(s)
      AF131800, BF529906, DA541045
      Consensus CDS
      CCDS74361.1
      UniProtKB/TrEMBL
      A0A0A0MRF5, E7EWX6
      Related
      ENSP00000351809.6, ENST00000358931.9
      Conserved Domains (1) summary
      cd01408
      Location:40228
      SIRT1; SIRT1: Eukaryotic group (class1) which includes human sirtuins SIRT1-3 and yeast Hst1-4; and are members of the SIR2 family of proteins, silent information regulator 2 (Sir2) enzymes which catalyze NAD+-dependent protein/histone deacetylation. Sir2 ...
    2. NM_012237.4NP_036369.2  NAD-dependent protein deacetylase sirtuin-2 isoform 1

      See identical proteins and their annotated locations for NP_036369.2

      Status: REVIEWED

      Description
      Transcript Variant: This variant (1) is the longest transcript and encodes the longest isoform (1).
      Source sequence(s)
      AF083107, BF529906
      Consensus CDS
      CCDS12523.1
      UniProtKB/Swiss-Prot
      A8K3V1, B2RB45, O95889, Q8IXJ6, Q924Y7, Q9P0G8, Q9UNT0, Q9Y6E9, U5TP13
      UniProtKB/TrEMBL
      A0AAG2T947
      Related
      ENSP00000249396.7, ENST00000249396.12
      Conserved Domains (1) summary
      cd01408
      Location:77331
      SIRT1; Eukaryotic group (class1) which includes human sirtuins SIRT1-3 and yeast Hst1-4; and are members of the SIR2 family of proteins, silent information regulator 2 (Sir2) enzymes which catalyze NAD+-dependent protein/histone deacetylation. Sir2 proteins ...
    3. NM_030593.3NP_085096.1  NAD-dependent protein deacetylase sirtuin-2 isoform 2

      See identical proteins and their annotated locations for NP_085096.1

      Status: REVIEWED

      Description
      Transcript Variant: This variant (2) lacks an exon in the 5' region, resulting in a downstream in-frame AUG start codon, as compared to variant 1. The resulting isoform (2) has a shorter N-terminus, as compared to isoform 1.
      Source sequence(s)
      BC003012
      Consensus CDS
      CCDS46069.1
      UniProtKB/TrEMBL
      A0AAG2T947
      Related
      ENSP00000375931.2, ENST00000392081.6
      Conserved Domains (2) summary
      cd01408
      Location:40294
      SIRT1; Eukaryotic group (class1) which includes human sirtuins SIRT1-3 and yeast Hst1-4; and are members of the SIR2 family of proteins, silent information regulator 2 (Sir2) enzymes which catalyze NAD+-dependent protein/histone deacetylation. Sir2 proteins ...
      pfam16602
      Location:313351
      USP19_linker; Linker region of USP19 deubiquitinase

    RNA

    1. NR_034146.1 RNA Sequence

      Status: REVIEWED

      Description
      Transcript Variant: This variant (4) lacks the 5' two exons but has an alternate 5' segment, resulting in an upstream open reading frame (uORF) with strong Kozak signal, as compared to variant 1. The uORF overlaps the downstream ORF (dORF), and inhibits the protein translation from the dORF. Therefore, this transcript is a nonsense-mediated mRNA decay candidate and can not make a functional protein.
      Source sequence(s)
      AK290716, BF346568, BF529906

    RefSeqs of Annotated Genomes: GCF_000001405.40-RS_2024_08

    The following sections contain reference sequences that belong to a specific genome build. Explain

    Reference GRCh38.p14 Primary Assembly

    Genomic

    1. NC_000019.10 Reference GRCh38.p14 Primary Assembly

      Range
      38878555..38899618 complement
      Download
      GenBank, FASTA, Sequence Viewer (Graphics)

    mRNA and Protein(s)

    1. XM_011526655.2XP_011524957.1  NAD-dependent protein deacetylase sirtuin-2 isoform X2

      See identical proteins and their annotated locations for XP_011524957.1

      UniProtKB/Swiss-Prot
      Q8IXJ6
      Conserved Domains (2) summary
      cd01408
      Location:7261
      SIRT1; SIRT1: Eukaryotic group (class1) which includes human sirtuins SIRT1-3 and yeast Hst1-4; and are members of the SIR2 family of proteins, silent information regulator 2 (Sir2) enzymes which catalyze NAD+-dependent protein/histone deacetylation. Sir2 ...
      pfam16602
      Location:280318
      USP19_linker; Linker region of USP19 deubiquitinase
    2. XM_047438468.1XP_047294424.1  NAD-dependent protein deacetylase sirtuin-2 isoform X1

      UniProtKB/TrEMBL
      A0AAG2T947
    3. XM_047438469.1XP_047294425.1  NAD-dependent protein deacetylase sirtuin-2 isoform X3

    Reference GRCh38.p14 PATCHES

    Genomic

    1. NW_014040929.1 Reference GRCh38.p14 PATCHES

      Range
      288227..309290 complement
      Download
      GenBank, FASTA, Sequence Viewer (Graphics)

    mRNA and Protein(s)

    1. XM_054331959.1XP_054187934.1  NAD-dependent protein deacetylase sirtuin-2 isoform X2

    2. XM_054331958.1XP_054187933.1  NAD-dependent protein deacetylase sirtuin-2 isoform X1

      UniProtKB/TrEMBL
      A0AAG2T947
    3. XM_054331960.1XP_054187935.1  NAD-dependent protein deacetylase sirtuin-2 isoform X3

    Alternate T2T-CHM13v2.0

    Genomic

    1. NC_060943.1 Alternate T2T-CHM13v2.0

      Range
      41682611..41703673 complement
      Download
      GenBank, FASTA, Sequence Viewer (Graphics)

    mRNA and Protein(s)

    1. XM_054320275.1XP_054176250.1  NAD-dependent protein deacetylase sirtuin-2 isoform X2

    2. XM_054320274.1XP_054176249.1  NAD-dependent protein deacetylase sirtuin-2 isoform X1

      UniProtKB/TrEMBL
      A0AAG2T947
    3. XM_054320276.1XP_054176251.1  NAD-dependent protein deacetylase sirtuin-2 isoform X3