FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\e(y)2
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General Information
Symbol
Dmel\e(y)2
Species
D. melanogaster
Name
enhancer of yellow 2
Annotation Symbol
CG15191
Feature Type
FlyBase ID
FBgn0000618
Gene Model Status
Stock Availability
Gene Summary
enhancer of yellow 2 (e(y)2) encodes a subunit of TREX2 mRNA nuclear export complex. It is also a subunit of histone deubiquitination module of SAGA complex. The product of e(y)2 is essential for general nuclear export of mRNA, transcription activation and gene positioning in the nuclei. [Date last reviewed: 2019-09-26] (FlyBase Gene Snapshot)
Also Known As

ENY2, late boundary complex, LBC

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
1-35
RefSeq locus
NC_004354 REGION:11576118..11576620
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (19 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:su(Hw); FB:FBgn0003567
inferred from direct assay
inferred from physical interaction with UniProtKB:Q7K2L1
inferred from physical interaction with UniProtKB:Q9Y1B2
inferred from physical interaction with UniProtKB:P08970
inferred from physical interaction with UniProtKB:Q9U3V9
inferred from physical interaction with UniProtKB:Q27272
inferred from physical interaction with FLYBASE:Taf9; FB:FBgn0000617
inferred from physical interaction with FLYBASE:Tbp; FB:FBgn0003687
inferred from physical interaction with FLYBASE:Taf10; FB:FBgn0028398
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000282153
inferred from biological aspect of ancestor with PANTHER:PTN000282153
inferred from electronic annotation with InterPro:IPR018783
Biological Process (6 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from direct assay
inferred from mutant phenotype
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
Cellular Component (7 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
located_in chromatin
inferred from direct assay
part_of DUBm complex
inferred from direct assay
inferred from direct assay
part_of nuclear pore
inferred from direct assay
located_in nucleus
inferred from direct assay
inferred from direct assay
part_of SAGA complex
inferred from direct assay
inferred from physical interaction with FLYBASE:Gcn5; FB:FBgn0020388
inferred from physical interaction with FLYBASE:Taf10; FB:FBgn0028398
inferred from physical interaction with FLYBASE:Ada2b; FB:FBgn0037555
inferred from physical interaction with FLYBASE:Nipped-A; FB:FBgn0053554
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
part_of DUBm complex
inferred from biological aspect of ancestor with PANTHER:PTN000282153
part_of SAGA complex
inferred from biological aspect of ancestor with PANTHER:PTN000282153
inferred from electronic annotation with InterPro:IPR018783
Protein Family (UniProt)
Belongs to the ENY2 family. (Q9VYX1)
Summaries
Gene Snapshot
enhancer of yellow 2 (e(y)2) encodes a subunit of TREX2 mRNA nuclear export complex. It is also a subunit of histone deubiquitination module of SAGA complex. The product of e(y)2 is essential for general nuclear export of mRNA, transcription activation and gene positioning in the nuclei. [Date last reviewed: 2019-09-26]
Gene Group (FlyBase)
TRANSCRIPTION EXPORT COMPLEX-2 -
TREX-2 is a multisubunit complex required for mRNA export through the nuclear pore. TREX-2 colocalizes with the nuclear pore complex at the nuclear periphery. (Adapted from FBrf0232523 and FBrf0200574).
SAGA COMPLEX -
The Spt-Ada-Gcn5 Acetyltransferase (SAGA) complex is a chromatin modifying complex. SAGA can acetylate and deubiquitinate histone residues. Also SAGA has been shown to regulate the transcriptional activation of a subset of genes. (Adapted from FBrf0228923, FBrf0219841 and FBrf0233349).
Protein Function (UniProtKB)
Involved in mRNA export coupled transcription activation by association with both the TREX-2/AMEX and the SAGA complexes (PubMed:18034162, PubMed:19947544, PubMed:27016737). The SAGA complex is a multiprotein complex that activates transcription by remodeling chromatin and mediating histone acetylation and deubiquitination (PubMed:11438676, PubMed:18206972). Within the SAGA complex, participates in a subcomplex that specifically deubiquitinates histone H2B (PubMed:18206972). The SAGA complex is recruited to specific gene promoters by activators, where it is required for transcription (PubMed:18034162, PubMed:19947544). Required for nuclear receptor-mediated transactivation (PubMed:20048002, PubMed:20714859). Involved in transcription elongation by recruiting the THO complex onto nascent mRNA (PubMed:20048002). The TREX-2/AMEX complex functions in docking export-competent ribonucleoprotein particles (mRNPs) to the nuclear entrance of the nuclear pore complex (nuclear basket) (PubMed:27016737). TREX-2/AMEX participates in mRNA export and accurate chromatin positioning in the nucleus by tethering genes to the nuclear periphery (PubMed:17643381, PubMed:27016737). Recruited to the su(Hw) insulators via its interaction with su(Hw) and participates in the barrier activity of such insulators (PubMed:17643381). In contrast, it does not participate in the enhancer-blocking activity of the su(Hw) insulators (PubMed:17643381).
(UniProt, Q9VYX1)
Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\e(y)2 for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q9VYX1)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Gene model reviewed during 5.40

Gene model reviewed during 5.45

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0073540
503
101
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0073385
11.5
101
4.42
Polypeptides with Identical Sequences

There is only one protein coding transcript and one polypeptide associated with this gene

Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

Component of the nuclear pore complex (NPC)-associated TREX-2/AMEX complex (anchoring and mRNA export complex), composed of e(y)2, xmas and PCID2 (PubMed:18034162, PubMed:27016737). Within the TREX-2/ AMEX complex, interactions with xmas is required for localization to the nuclear periphery (PubMed:18034162). Component of the SAGA transcription coactivator-HAT complexes, at least composed of Ada2b, e(y)2, Pcaf/Gcn5, Taf10 and Nipped-A/Trrap (PubMed:18034162, PubMed:19947544). Within the SAGA complex, e(y)2, Sgf11, and not/nonstop form an additional subcomplex of SAGA called the DUB module (deubiquitination module) (By similarity). Component of the THO complex, composed of at least e(y)2, HPR1, THO2, THOC5, THOC6 and THOC7 (PubMed:20048002, PubMed:20714859). Interacts with Taf9 (PubMed:11438676). Interacts with su(Hw) (via zinc fingers) (PubMed:17643381). Interacts with the nuclear pore complex (NPC) (PubMed:18034162, PubMed:20048002). Interaction between the TREX-2/AMEX complex and the ORC complex is required for ORC localization to mRNPs, and consequently mRNA export (PubMed:27016737, Ref.11). Within the TREX-2/AMEX-ORC complex, interacts with Orc6 and (via N-terminus or C-terminus) with Orc3 (PubMed:27016737, Ref.11). Interacts with the zinc finger protein CG9890 (PubMed:30713769).

(UniProt, Q9VYX1)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\e(y)2 using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

1.57

Transcript Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in chromatin
inferred from direct assay
part_of DUBm complex
inferred from direct assay
inferred from direct assay
part_of nuclear pore
inferred from direct assay
located_in nucleus
inferred from direct assay
inferred from direct assay
part_of SAGA complex
inferred from direct assay
inferred from physical interaction with FLYBASE:Gcn5; FB:FBgn0020388
inferred from physical interaction with FLYBASE:Taf10; FB:FBgn0028398
inferred from physical interaction with FLYBASE:Ada2b; FB:FBgn0037555
inferred from physical interaction with FLYBASE:Nipped-A; FB:FBgn0053554
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\e(y)2 in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 3 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 7 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of e(y)2
Transgenic constructs containing regulatory region of e(y)2
Aberrations (Deficiencies and Duplications) ( 0 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (1)
13 of 14
Yes
Yes
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (1)
13 of 14
Yes
Yes
Mus musculus (laboratory mouse) (1)
12 of 14
Yes
Yes
Xenopus tropicalis (Western clawed frog) (1)
10 of 13
Yes
Yes
Danio rerio (Zebrafish) (1)
12 of 14
Yes
Yes
Caenorhabditis elegans (Nematode, roundworm) (0)
Anopheles gambiae (African malaria mosquito) (0)
Arabidopsis thaliana (thale-cress) (1)
12 of 13
Yes
Yes
Saccharomyces cerevisiae (Brewer's yeast) (1)
9 of 13
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (1)
6 of 12
Yes
Yes
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:e(y)2. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (1)
9 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 1 )
    Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    Functional Complementation Data
    Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
    Interactions
    Summary of Physical Interactions
    Interaction Browsers

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    RNA-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    Interaction Browsers

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    enhanceable
    External Data
    Subunit Structure (UniProtKB)
    Component of the nuclear pore complex (NPC)-associated TREX-2/AMEX complex (anchoring and mRNA export complex), composed of e(y)2, xmas and PCID2 (PubMed:18034162, PubMed:27016737). Within the TREX-2/ AMEX complex, interactions with xmas is required for localization to the nuclear periphery (PubMed:18034162). Component of the SAGA transcription coactivator-HAT complexes, at least composed of Ada2b, e(y)2, Pcaf/Gcn5, Taf10 and Nipped-A/Trrap (PubMed:18034162, PubMed:19947544). Within the SAGA complex, e(y)2, Sgf11, and not/nonstop form an additional subcomplex of SAGA called the DUB module (deubiquitination module) (By similarity). Component of the THO complex, composed of at least e(y)2, HPR1, THO2, THOC5, THOC6 and THOC7 (PubMed:20048002, PubMed:20714859). Interacts with Taf9 (PubMed:11438676). Interacts with su(Hw) (via zinc fingers) (PubMed:17643381). Interacts with the nuclear pore complex (NPC) (PubMed:18034162, PubMed:20048002). Interaction between the TREX-2/AMEX complex and the ORC complex is required for ORC localization to mRNPs, and consequently mRNA export (PubMed:27016737, Ref.11). Within the TREX-2/AMEX-ORC complex, interacts with Orc6 and (via N-terminus or C-terminus) with Orc3 (PubMed:27016737, Ref.11). Interacts with the zinc finger protein CG9890 (PubMed:30713769).
    (UniProt, Q9VYX1 )
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    FlyBase
    External Links
    External Data
    Linkouts
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    X
    Recombination map
    1-35
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    10C7-10C7
    Limits computationally determined from genome sequence between P{EP}CG32666EP1452 and P{EP}Ptp10DEP1172
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    10C-10C
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (5)
    Genomic Clones (6)
     

    Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

    cDNA Clones (6)
     

    Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see JBrowse for alignment of the cDNAs and ESTs to the gene model.

    cDNA clones, fully sequenced
    BDGP DGC clones
    Other clones
    Drosophila Genomics Resource Center cDNA clones

    For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

    cDNA Clones, End Sequenced (ESTs)
    BDGP DGC clones
      RNAi and Array Information
      Linkouts
      DRSC - Results frm RNAi screens
      Antibody Information
      Laboratory Generated Antibodies
      Commercially Available Antibodies
       
      Cell Line Information
      Publicly Available Cell Lines
       
        Other Stable Cell Lines
         
          Other Comments

          The effect of mutations in the e(y)2 locus on mutagenesis in the y2ns scme double superunstable system has been analysed.

          Mutations at the e(y)2 loci enhance the phenotypic expression of several insertion mutations of y without exerting any effect on mdg4 induced mutations at other loci. One mutation in e(y)2 is due to insertion of a Stalker element.

          Modifies the expression of y2, a gypsy-induced allele, but not other gypsy-induced mutations. The effects of all e(y) alleles on bristle color persist in the presence of su(Hw)2. No effect on the phenotype of y+.

          Relationship to Other Genes
          Source for database merge of

          Source for merge of: e(y)2 CG15191

          Additional comments

          The e(y)2 gene may have been derived from the e(y)2b gene by retroposition.

          "e(y)2" appears to have been derived from "e(y)2b" by retroposition; "e(y)2"is composed of a single exon and contains 17bp direct repeats beginning at positions -12 and +528 nucleotides relative to the transcription start site (these repeats are a hallmark of retroposition).

          FlyBase curator comment: the insertion in the "e03298" Exelixis line (PBac{RB}CG11695e03298) was originally assigned to the e(y)2 gene in FBrf0179797, resulting in the "e(y)2e03298" (FBal0159294) allele. However, FBrf0184340 shows that the insertion is actually within CG11695.

          Source for merge of e(y)2 CG15191 was sequence comparison ( date:000815 ).

          Nomenclature History
          Source for database identify of
          Nomenclature comments
          Etymology
          Synonyms and Secondary IDs (13)
          Reported As
          Symbol Synonym
          Name Synonyms
          enhancer of yellow 2
          late boundary complex
          Secondary FlyBase IDs
          • FBgn0030315
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 31 )
          Sequence Crossreferences
          NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
          GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
          GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
          RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
          UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
          UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
          Other crossreferences
          AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
          BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
          DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
          EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
          FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
          FlyMine - An integrated database for Drosophila genomics
          InterPro - A database of protein families, domains and functional sites
          KEGG Genes - Molecular building blocks of life in the genomic space.
          MARRVEL_MODEL - MARRVEL (model organism gene)
          Linkouts
          BioGRID - A database of protein and genetic interactions.
          Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
          DroID - A comprehensive database of gene and protein interactions.
          DRSC - Results frm RNAi screens
          Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
          FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
          FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
          Flygut - An atlas of the Drosophila adult midgut
          iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
          MIST (genetic) - An integrated Molecular Interaction Database
          MIST (protein-protein) - An integrated Molecular Interaction Database
          References (99)