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    GOLGA3 golgin A3 [ Homo sapiens (human) ]

    Gene ID: 2802, updated on 10-Dec-2024

    Summary

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    Official Symbol
    GOLGA3provided by HGNC
    Official Full Name
    golgin A3provided by HGNC
    Primary source
    HGNC:HGNC:4426
    See related
    Ensembl:ENSG00000090615 MIM:602581; AllianceGenome:HGNC:4426
    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
    MEA-2; GCP170
    Summary
    The Golgi apparatus, which participates in glycosylation and transport of proteins and lipids in the secretory pathway, consists of a series of stacked cisternae (flattened membrane sacs). Interactions between the Golgi and microtubules are thought to be important for the reorganization of the Golgi after it fragments during mitosis. This gene encodes a member of the golgin family of proteins which are localized to the Golgi. Its encoded protein has been postulated to play a role in nuclear transport and Golgi apparatus localization. Several alternatively spliced transcript variants that encode different protein isoforms have been described for this gene. [provided by RefSeq, Feb 2010]
    Expression
    Ubiquitous expression in colon (RPKM 8.3), brain (RPKM 6.7) and 25 other tissues See more
    Orthologs
    mouse all
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    Genomic context

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    See GOLGA3 in Genome Data Viewer
    Location:
    12q24.33
    Exon count:
    30
    Annotation release Status Assembly Chr Location
    RS_2024_08 current GRCh38.p14 (GCF_000001405.40) 12 NC_000012.12 (132768914..132829081, complement)
    RS_2024_08 current T2T-CHM13v2.0 (GCF_009914755.1) 12 NC_060936.1 (132819976..132879866, complement)
    RS_2024_09 previous assembly GRCh37.p13 (GCF_000001405.25) 12 NC_000012.11 (133345500..133405667, complement)

    Chromosome 12 - NC_000012.12Genomic Context describing neighboring genes Neighboring gene RNA, 5S ribosomal pseudogene 379 Neighboring gene ankyrin repeat and LEM domain containing 2 Neighboring gene TPD52 like 2 pseudogene Neighboring gene ReSE screen-validated silencer GRCh37_chr12:133334029-133334208 Neighboring gene H3K27ac hESC enhancer GRCh37_chr12:133337764-133338632 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 7387 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 7388 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 7389 Neighboring gene Neanderthal introgressed variant-containing enhancer experimental_26051 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 7390 Neighboring gene H3K4me1 hESC enhancer GRCh37_chr12:133365291-133365792 Neighboring gene Neanderthal introgressed variant-containing enhancer experimental_26074 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 7391 Neighboring gene ATAC-STARR-seq lymphoblastoid silent region 5128 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 7392 Neighboring gene ATAC-STARR-seq lymphoblastoid silent region 5129 Neighboring gene ATAC-STARR-seq lymphoblastoid silent region 5130 Neighboring gene BRD4-independent group 4 enhancer GRCh37_chr12:133412143-133413342 Neighboring gene ribosomal protein S11 pseudogene 5 Neighboring gene RNA, U6 small nuclear 327, pseudogene Neighboring gene checkpoint with forkhead and ring finger domains Neighboring gene H3K4me1 hESC enhancer GRCh37_chr12:133428785-133429286 Neighboring gene H3K4me1 hESC enhancer GRCh37_chr12:133429287-133429786 Neighboring gene BRD4-independent group 4 enhancer GRCh37_chr12:133433141-133434340 Neighboring gene H3K4me1 hESC enhancer GRCh37_chr12:133449077-133449774 Neighboring gene H3K4me1 hESC enhancer GRCh37_chr12:133449775-133450470 Neighboring gene H3K27ac-H3K4me1 hESC enhancer GRCh37_chr12:133450471-133451166 Neighboring gene H3K27ac-H3K4me1 hESC enhancer GRCh37_chr12:133451167-133451862 Neighboring gene ATAC-STARR-seq lymphoblastoid silent region 5131 Neighboring gene ATAC-STARR-seq lymphoblastoid active region 7393 Neighboring gene OCT4-NANOG-H3K27ac-H3K4me1 hESC enhancer GRCh37_chr12:133464984-133465786 Neighboring gene CHFR divergent transcript

    Genomic regions, transcripts, and products

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    Go to nucleotide: Graphics FASTA GenBank

    Expression

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    • Project title: Tissue-specific circular RNA induction during human fetal development
    • Description: 35 human fetal samples from 6 tissues (3 - 7 replicates per tissue) collected between 10 and 20 weeks gestational time were sequenced using Illumina TruSeq Stranded Total RNA
    • BioProject: PRJNA270632
    • Publication: PMID 26076956
    • Analysis date: Mon Apr 2 22:54:59 2018

    Bibliography

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    Related articles in PubMed

    1. Three basic residues of intracellular loop 3 of the beta-1 adrenergic receptor are required for golgin-160-dependent trafficking. Gilbert CE, et al. Int J Mol Sci, 2014 Feb 20. PMID 24566136, Free PMC Article
    2. Golgin160 recruits the dynein motor to position the Golgi apparatus. Yadav S, et al. Dev Cell, 2012 Jul 17. PMID 22814606, Free PMC Article
    3. Molecular characterization of GCP170, a 170-kDa protein associated with the cytoplasmic face of the Golgi membrane. Misumi Y, et al. J Biol Chem, 1997 Sep 19. PMID 9295333
    4. Identification of a common autoantigenic epitope of protein disulfide isomerase, golgin-160 and voltage-gated potassium channel in type 1 diabetes. Fierabracci A, et al. Diabetes Res Clin Pract, 2010 May. PMID 20170975
    5. Caspase-resistant Golgin-160 disrupts apoptosis induced by secretory pathway stress and ligation of death receptors. Maag RS, et al. Mol Biol Cell, 2005 Jun. PMID 15829563, Free PMC Article

    See all (105) citations in PubMed

    GeneRIFs: Gene References Into Functions

    What's a GeneRIF?
    1. An exome-first approach to aid in the diagnosis of primary ciliary dyskinesia.
      Title: An exome-first approach to aid in the diagnosis of primary ciliary dyskinesia.
    2. Golgin-160 disperses from Golgi membranes in cells subjected to cold temperature. Activated ARF1 can prevent cold-induced dispersal of golgin-160.
      Title: Commonly used trafficking blocks disrupt ARF1 activation and the localization and function of specific Golgi proteins.
    3. The golgin-160 binds directly to the third intracellular loop of beta1AR and that this binding depends on three basic residues in this loop.
      Title: Three basic residues of intracellular loop 3 of the beta-1 adrenergic receptor are required for golgin-160-dependent trafficking.
    4. Golgi protein golgin160 recruits dynein to Golgi membranes.
      Title: Golgin160 recruits the dynein motor to position the Golgi apparatus.
    5. A common epitope of proteins golgin-160, voltage-gated potassium channel and disulfide isomerase was identified by screening with autoantibodies of a type 1 diabetic patient
      Title: Identification of a common autoantigenic epitope of protein disulfide isomerase, golgin-160 and voltage-gated potassium channel in type 1 diabetes.
    6. Clinical trial of gene-disease association and gene-environment interaction. (HuGE Navigator)
      Title: Personalized smoking cessation: interactions between nicotine dose, dependence and quit-success genotype score.
    7. nuclear translocation of golgin-160-(140-311) is a highly coordinated event regulated not only by cleavage of the golgin-160 head but also by the oxidation state of GCP60
      Title: Identification of a redox-sensitive cysteine in GCP60 that regulates its interaction with golgin-160.
    8. results indicate that GCP16 is the acylated membrane protein, associated with GCP170, and possibly involved in vesicular transport from the Golgi to the cell surface
      Title: Identification and characterization of GCP16, a novel acylated Golgi protein that interacts with GCP170.
    9. NH2 terminal domain of Golgin 160 has a role in nuclear transport and Golgi apparatus localization
      Title: The NH2-terminal domain of Golgin-160 contains both Golgi and nuclear targeting information.
    10. golgin-160 may promote efficient surface delivery of a subset of cargo molecules
      Title: Golgin-160 promotes cell surface expression of the beta-1 adrenergic receptor.
    Submit: New GeneRIF Correction See all GeneRIFs (13)

    Phenotypes

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    Find tests for this gene in the NIH Genetic Testing Registry (GTR)

    Review eQTL and phenotype association data in this region using PheGenI

    EBI GWAS Catalog

    Description
    Genome-wide association study of antibody response to smallpox vaccine.
    EBI GWAS Catalog

    Variation

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    See variants in ClinVar

    See studies and variants in dbVar

    See Variation Viewer (GRCh37.p13)

    See Variation Viewer (GRCh38)

    Interactions

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    Products Interactant Other Gene Complex Source Pubs Description

    General gene information

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    Markers

    Homology

    Gene Ontology Provided by GOA

    Function Evidence Code Pubs
    enables cadherin binding HDA PubMed 
    enables protein binding IPI
    Inferred from Physical Interaction
    more info
    		 PubMed 
    Component Evidence Code Pubs
    located_in Golgi apparatus IDA
    Inferred from Direct Assay
    more info
    		  
    located_in Golgi apparatus TAS
    Traceable Author Statement
    more info
    		 PubMed 
    located_in Golgi cisterna membrane IEA
    Inferred from Electronic Annotation
    more info
    		  
    located_in Golgi membrane IDA
    Inferred from Direct Assay
    more info
    		 PubMed 
    located_in cytosol IDA
    Inferred from Direct Assay
    more info
    		 PubMed 
    located_in membrane HDA PubMed 
    located_in nucleolus IDA
    Inferred from Direct Assay
    more info
    		  
    located_in nucleoplasm IDA
    Inferred from Direct Assay
    more info
    		  

    General protein information

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    Preferred Names
    golgin subfamily A member 3
    Names
    Golgi membrane associated protein
    Golgi peripheral membrane protein
    SY2/SY10 protein
    golgi autoantigen, golgin subfamily a, 3
    golgi complex-associated protein of 170 kDa
    golgin-160
    golgin-165
    male enhanced antigen-2

    NCBI Reference Sequences (RefSeq)

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    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.

    Genomic

    1. NG_029887.1 RefSeqGene

      Range
      4972..64927
      Download
      GenBank, FASTA, Sequence Viewer (Graphics)

    mRNA and Protein(s)

    1. NM_001172557.2 → NP_001166028.1  golgin subfamily A member 3 isoform 2

      See identical proteins and their annotated locations for NP_001166028.1

      Status: REVIEWED

      Description
      Transcript Variant: This variant (2) uses a different segment for its 5' UTR and contains a different 3' end, compared to variant 1. The resulting protein (isoform 2) has a shorter and distinct C-terminus when it is compared to isoform 1.
      Source sequence(s)
      AC127070, BC146675, D63997
      Consensus CDS
      CCDS53846.1
      UniProtKB/TrEMBL
      A0A8I5KVX9
      Related
      ENSP00000442603.1, ENST00000545875.4
      Conserved Domains (2) summary
      pfam05010
      Location:812 → 997
      TACC; Transforming acidic coiled-coil-containing protein (TACC)
      cl19219
      Location:812 → 886
      DUF342; Protein of unknown function (DUF342)

    2. NM_001389683.1 → NP_001376612.1  golgin subfamily A member 3 isoform 1

      Status: REVIEWED

      Source sequence(s)
      AC127070, AC136467
      Consensus CDS
      CCDS9281.1
      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0
      Related
      ENSP00000410378.2, ENST00000450791.8
      Conserved Domains (2) summary
      COG1196
      Location:630 → 1358
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]
      TIGR02168
      Location:393 → 743
      SMC_prok_B; chromosome segregation protein SMC, common bacterial type

    3. NM_001389684.1 → NP_001376613.1  golgin subfamily A member 3 isoform 1

      Status: REVIEWED

      Source sequence(s)
      AC127070, AC136467
      Consensus CDS
      CCDS9281.1
      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0
      Conserved Domains (2) summary
      COG1196
      Location:630 → 1358
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]
      TIGR02168
      Location:393 → 743
      SMC_prok_B; chromosome segregation protein SMC, common bacterial type

    4. NM_001389685.1 → NP_001376614.1  golgin subfamily A member 3 isoform 1

      Status: REVIEWED

      Source sequence(s)
      AC127070, AC136467
      Consensus CDS
      CCDS9281.1
      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0
      Conserved Domains (2) summary
      COG1196
      Location:630 → 1358
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]
      TIGR02168
      Location:393 → 743
      SMC_prok_B; chromosome segregation protein SMC, common bacterial type

    5. NM_001389686.1 → NP_001376615.1  golgin subfamily A member 3 isoform 3

      Status: REVIEWED

      Source sequence(s)
      AC127070, AC136467
      Consensus CDS
      CCDS91785.1
      UniProtKB/TrEMBL
      A0A8I5KSF0, A0A8I5KWY9
      Related
      ENSP00000510765.1, ENST00000688114.1
      Conserved Domains (2) summary
      COG1196
      Location:590 → 1318
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]
      TIGR02168
      Location:353 → 703
      SMC_prok_B; chromosome segregation protein SMC, common bacterial type

    6. NM_001389687.1 → NP_001376616.1  golgin subfamily A member 3 isoform 3

      Status: REVIEWED

      Source sequence(s)
      AC127070, AC136467
      Consensus CDS
      CCDS91785.1
      UniProtKB/TrEMBL
      A0A8I5KSF0, A0A8I5KWY9
      Conserved Domains (2) summary
      COG1196
      Location:590 → 1318
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]
      TIGR02168
      Location:353 → 703
      SMC_prok_B; chromosome segregation protein SMC, common bacterial type

    7. NM_001389688.1 → NP_001376617.1  golgin subfamily A member 3 isoform 3

      Status: REVIEWED

      Source sequence(s)
      AC127070, AC136467
      Consensus CDS
      CCDS91785.1
      UniProtKB/TrEMBL
      A0A8I5KSF0, A0A8I5KWY9
      Related
      ENSP00000509869.1, ENST00000691696.1
      Conserved Domains (2) summary
      COG1196
      Location:590 → 1318
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]
      TIGR02168
      Location:353 → 703
      SMC_prok_B; chromosome segregation protein SMC, common bacterial type

    8. NM_001389689.1 → NP_001376618.1  golgin subfamily A member 3 isoform 4

      Status: REVIEWED

      Source sequence(s)
      AC127070, AC136467
      UniProtKB/TrEMBL
      A0A8I5KSF0
      Conserved Domains (1) summary
      COG1196
      Location:539 → 1267
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]

    9. NM_005895.4 → NP_005886.2  golgin subfamily A member 3 isoform 1

      See identical proteins and their annotated locations for NP_005886.2

      Status: REVIEWED

      Description
      Transcript Variant: This variant (1) represents the longer transcript and it encodes the longer protein (isoform 1).
      Source sequence(s)
      AC127070, AC136467, AF485338, BC060826, BF902971, D63997
      Consensus CDS
      CCDS9281.1
      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0
      Related
      ENSP00000204726.3, ENST00000204726.9
      Conserved Domains (2) summary
      COG1196
      Location:630 → 1358
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]
      TIGR02168
      Location:393 → 743
      SMC_prok_B; chromosome segregation protein SMC, common bacterial type

    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_000012.12 Reference GRCh38.p14 Primary Assembly

      Range
      132768914..132829081 complement
      Download
      GenBank, FASTA, Sequence Viewer (Graphics)

    mRNA and Protein(s)

    1. XM_047428706.1 → XP_047284662.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    2. XM_047428708.1 → XP_047284664.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    3. XM_024448941.2 → XP_024304709.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0
      Conserved Domains (2) summary
      COG1196
      Location:630 → 1358
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]
      TIGR02168
      Location:393 → 743
      SMC_prok_B; chromosome segregation protein SMC, common bacterial type

    4. XM_047428707.1 → XP_047284663.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    5. XM_024448940.2 → XP_024304708.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0
      Conserved Domains (2) summary
      COG1196
      Location:630 → 1358
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]
      TIGR02168
      Location:393 → 743
      SMC_prok_B; chromosome segregation protein SMC, common bacterial type

    6. XM_047428705.1 → XP_047284661.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    7. XM_047428709.1 → XP_047284665.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    8. XM_005266167.5 → XP_005266224.1  golgin subfamily A member 3 isoform X2

      UniProtKB/TrEMBL
      A0A8I5KSF0
      Related
      ENSP00000508770.1, ENST00000690511.1
      Conserved Domains (2) summary
      COG1196
      Location:439 → 1167
      Smc; Chromosome segregation ATPase [Cell cycle control, cell division, chromosome partitioning]
      cl19219
      Location:621 → 695
      DUF342; Protein of unknown function (DUF342)

    9. XM_011534794.3 → XP_011533096.1  golgin subfamily A member 3 isoform X3

      Conserved Domains (3) summary
      TIGR02168
      Location:20 → 694
      SMC_prok_B; chromosome segregation protein SMC, common bacterial type
      pfam01496
      Location:446 → 526
      V_ATPase_I; V-type ATPase 116kDa subunit family
      cl19219
      Location:164 → 238
      DUF342; Protein of unknown function (DUF342)

    Alternate T2T-CHM13v2.0

    Genomic

    1. NC_060936.1 Alternate T2T-CHM13v2.0

      Range
      132819976..132879866 complement
      Download
      GenBank, FASTA, Sequence Viewer (Graphics)

    mRNA and Protein(s)

    1. XM_054371760.1 → XP_054227735.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    2. XM_054371762.1 → XP_054227737.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    3. XM_054371759.1 → XP_054227734.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    4. XM_054371761.1 → XP_054227736.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    5. XM_054371758.1 → XP_054227733.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    6. XM_054371757.1 → XP_054227732.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    7. XM_054371756.1 → XP_054227731.1  golgin subfamily A member 3 isoform X1

      UniProtKB/Swiss-Prot
      A5PKX6, O43241, Q08378, Q6P9C7, Q86XW3, Q8TDA9, Q8WZA3
      UniProtKB/TrEMBL
      A0A8I5KSF0

    8. XM_054371763.1 → XP_054227738.1  golgin subfamily A member 3 isoform X2

      UniProtKB/TrEMBL
      A0A8I5KSF0

    9. XM_054371764.1 → XP_054227739.1  golgin subfamily A member 3 isoform X3

    Nucleotide Protein
    Heading Accession and Version
    Protein Accession Links
    GenPept Link UniProtKB Link
    Q08378.2 GenPept UniProtKB/Swiss-Prot:Q08378

    Gene LinkOut

    The following LinkOut resources are supplied by external providers. These providers are responsible for maintaining the links.

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