FB2025_02 , released April 17, 2025
Gene: Dmel\asun
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General Information
Symbol
Dmel\asun
Species
D. melanogaster
Name
asunder
Annotation Symbol
CG6814
Feature Type
protein_coding_gene
FlyBase ID
FBgn0020407
Gene Model Status
Current
Stock Availability
7 publicly available
Gene Summary
Component of the Integrator complex, a complex involved in the transcription of small nuclear RNAs (snRNA) and their 3'-box-dependent processing (PubMed:23097424). Involved in the 3'-end processing of the U7 snRNA, and also the spliceosomal snRNAs U1 and U5 (PubMed:23097424). Plays a role as a regulator of spermatogenesis (PubMed:19357193). Crucial regulator of the mitotic cell cycle and development (PubMed:15737938, PubMed:19357193). Required for the correct dynein-dynactin perinuclear localization important for nucleus-centrosome coupling that occur upon meiotic progression of primary spermatocytes (PubMed:19357193, PubMed:23904267). Plays a role in sperm motility and fertility (PubMed:19357193). May have a role in the PNG/PLU/GNU pathway (PubMed:15737938). (UniProt, Q9VEX5)
Contribute a Gene Snapshot for this gene.
All Summaries
Also Known As

Mat89Bb, IntS13

Key Links
Genomic Location
Cytogenetic map
89B9-89B9
Sequence location
Recombination map
3-58
RefSeq locus
NT_033777 REGION:16264271..16266891
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (14 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (10 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
involved_in centrosome localization
inferred from mutant phenotype
(Sitaram et al., 2014, Sitaram et al., 2012)
involved_in dorsal appendage formation
inferred from mutant phenotype
(Sitaram et al., 2014)
involved_in flagellated sperm motility
inferred from mutant phenotype
(Anderson et al., 2009)
involved_in oocyte nucleus localization involved in oocyte dorsal/ventral axis specification
inferred from mutant phenotype
(Sitaram et al., 2014)
involved_in posterior mRNA localization involved in anterior/posterior axis specification
inferred from mutant phenotype
(Sitaram et al., 2014)
involved_in regulation of fertilization
inferred from mutant phenotype
(Anderson et al., 2009)
involved_in regulation of mitotic cell cycle
inferred from mutant phenotype
(Lee et al., 2005)
involved_in RNA polymerase II transcription initiation surveillance
inferred from direct assay
(Fujiwara et al., 2023, Huang et al., 2020, Tatomer et al., 2019)
involved_in snRNA 3'-end processing
inferred from direct assay
(Chen et al., 2012)
involved_in spermatogenesis
inferred from mutant phenotype
(Sitaram et al., 2012, Anderson et al., 2009)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
involved_in centrosome localization
inferred from biological aspect of ancestor with PANTHER:PTN000986954
(GO Reference Genome Project, 2011-)
involved_in regulation of mitotic cell cycle
inferred from biological aspect of ancestor with PANTHER:PTN000986954
(GO Reference Genome Project, 2011-)
Cellular Component (4 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
(Jodoin et al., 2013)
inferred from direct assay
(Chen et al., 2012, Anderson et al., 2009)
part_of INTAC complex
inferred from direct assay
(Fujiwara et al., 2023, Huang et al., 2020)
part_of integrator complex
inferred from direct assay
(Lin et al., 2024, Chen et al., 2012)
inferred from direct assay
(Tatomer et al., 2019)
is_active_in nucleus
inferred from direct assay
(Lin et al., 2024)
located_in nucleus
inferred from direct assay
(Jodoin et al., 2013, Chen et al., 2012, Anderson et al., 2009)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
part_of integrator complex
inferred from biological aspect of ancestor with PANTHER:PTN000986954
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the asunder family. (Q9VEX5)
Protein Signatures (InterPro)
Summaries
Gene Group (FlyBase)
INTEGRATOR COMPLEX -
The Integrator complex is an RNA Polymerase II (Pol II)-associated endonucleolytic complex that mediates 3'-end generation of small nuclear RNAs (snRNAs) and other non-coding RNAs, and modulates Pol II promoter-proximal pausing and transcript elongation. (Adapted from PMID:36180603.)
INTEGRATOR-PP2A COMPLEX -
The Integrator-PP2A complex (INTAC) is a complex formed between the endonucleolytic integrator complex and a serine/threonine-protein phosphatase 2A (PP2A) module. The PP2A module is formed from a PP2A-A scaffold subunit and a PP2A-C catalytic subunit, but lacks the PP2A-B regulatory subunit characteristic of canonical PP2A complexes. INTAC inhibits promoter-proximal pause release of RNA polymerase II (Pol II) by the dephosphorylation of a number of targets including the C-terminal domain of Pol II. (Adapted from PMID:36180603.)
Protein Function (UniProtKB)
Component of the Integrator complex, a complex involved in the transcription of small nuclear RNAs (snRNA) and their 3'-box-dependent processing (PubMed:23097424). Involved in the 3'-end processing of the U7 snRNA, and also the spliceosomal snRNAs U1 and U5 (PubMed:23097424). Plays a role as a regulator of spermatogenesis (PubMed:19357193). Crucial regulator of the mitotic cell cycle and development (PubMed:15737938, PubMed:19357193). Required for the correct dynein-dynactin perinuclear localization important for nucleus-centrosome coupling that occur upon meiotic progression of primary spermatocytes (PubMed:19357193, PubMed:23904267). Plays a role in sperm motility and fertility (PubMed:19357193). May have a role in the PNG/PLU/GNU pathway (PubMed:15737938).
(UniProt, Q9VEX5)
Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\asun 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 Q9VEX5)

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 5.47

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
asun-RA
FBtr0083259
NM_079655
2497
689
Additional Transcript Data and Comments
Reported size (kB)

2.5 (northern blot)

(Stebbings et al., 1998)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
asun-PA
FBpp0082713
75.7
689
8.57
Q9VEX5
NP_524379
AAF55290
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)

545 (aa); 60 (kD predicted)

(Stebbings et al., 1998)
Comments
External Data
Subunit Structure (UniProtKB)

Belongs to the multiprotein complex Integrator, at least composed of IntS1, IntS2, IntS3, IntS4, omd/IntS5, IntS6, defl/IntS7, IntS8, IntS9, IntS10, IntS11, IntS12, asun/IntS13 and IntS14.

(UniProt, Q9VEX5)
Post Translational Modification

Phosphorylated.

(UniProt, Q9VEX5)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
UniProt
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\asun 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).

0.06

Transcript Expression
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 9

Comment: reference states 0-4 hr AEL

(Stebbings et al., 1998)
adult stage
testis
(Stebbings et al., 1998)
ovary
(Stebbings et al., 1998)
adult stageoogenesis stage S9 -- S10
nurse cell
(Stebbings et al., 1998)
adult stage & oogenesis stage S10
follicle cell
(Stebbings et al., 1998)
adult stageoogenesis stage S10 -- S14
oocyte
(Stebbings et al., 1998)
Additional Descriptive Data

The 2.5 kb Mat89Bb transcript is expressed at high levels in ovaries and in 0-4 hour embryos, and at low levels in other stages and tissues, including testes. Mat89Bb transcripts are detected in nurse cells starting at stage S9, and are exported to the oocyte at stage S10. At stage S10, expression is also detected in follicle cells.

(Stebbings et al., 1998)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
(Jodoin et al., 2013)
inferred from direct assay
(Chen et al., 2012, Anderson et al., 2009)
part_of INTAC complex
inferred from direct assay
(Fujiwara et al., 2023, Huang et al., 2020)
part_of integrator complex
inferred from direct assay
(Lin et al., 2024, Chen et al., 2012)
inferred from direct assay
(Tatomer et al., 2019)
is_active_in nucleus
inferred from direct assay
(Lin et al., 2024)
located_in nucleus
inferred from direct assay
(Jodoin et al., 2013, Chen et al., 2012, Anderson et al., 2009)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\asun in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
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
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
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 2 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 9 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of asun
Transgenic constructs containing regulatory region of asun
Aberrations (Deficiencies and Duplications) ( 2 )
Inferred from experimentation ( 2 )
Gene disrupted in
Df(3R)Exel6268
(Ryder, 2004.4.26)
Df(3R)Exel7329
(Ryder, 2004.4.26)
Inferred from location ( 7 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (1)
Hsap\INTS13
14 of 14
Yes
Yes
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (1)
Rnor\Ints13
14 of 14
Yes
Yes
Mus musculus (laboratory mouse) (1)
Mmus\Ints13
13 of 14
Yes
Yes
Xenopus tropicalis (Western clawed frog) (2)
Xtro\ints13
11 of 13
Yes
Yes
Xtro\LOC100497319
1 of 13
No
No
Danio rerio (Zebrafish) (1)
Drer\ints13
14 of 14
Yes
Yes
Caenorhabditis elegans (Nematode, roundworm) (1)
Cele\ints-13
11 of 14
Yes
Yes
Anopheles gambiae (African malaria mosquito) (1)
Agam\AgaP_AGAP002295
12 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:asun. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
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 ( 0 )
    Allele
    Disease
    Interaction
    References
    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?
    INTS13; integrator complex subunit 13
    14 of 14
    615079
        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
        Summary of Genetic Interactions
        Interaction Browsers
        View in FlyBase Interactions Browser   View in MIST tool (external link)
        Please look at the allele data for full details of the genetic interactions
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        asun
        enhanceable
        Dhc64C
        (Anderson et al., 2009)
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        External Data
        Subunit Structure (UniProtKB)
        Belongs to the multiprotein complex Integrator, at least composed of IntS1, IntS2, IntS3, IntS4, omd/IntS5, IntS6, defl/IntS7, IntS8, IntS9, IntS10, IntS11, IntS12, asun/IntS13 and IntS14.
        (UniProt, Q9VEX5 )
        Linkouts
        BioGRID - A database of protein and genetic interactions.
        DroID - A comprehensive database of gene and protein interactions.
        MIST (protein-protein) - An integrated Molecular Interaction Database
        Pathways
        Signaling Pathways (FlyBase)
        Metabolic Pathways
        FlyBase
        External Links
        External Data
        Linkouts
        Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
        Class of Gene
        Genomic Location and Detailed Mapping Data
        Chromosome (arm)
        3R
        Recombination map
        3-58
        Cytogenetic map
        89B9-89B9
        Sequence location
        FlyBase Computed Cytological Location
        Cytogenetic map
        Evidence for location
        89B9-89B9
        Limits computationally determined from genome sequence between P{PZ}blp01618 and P{PZ}gish04895&P{EP}EP3171
        Experimentally Determined Cytological Location
        Cytogenetic map
        Notes
        References
        89B-89B
        (determined by in situ hybridisation)
        (Stebbings et al., 1998)
        Experimentally Determined Recombination Data
        Location

        3-58

        (FlyBase, 2016)

        3-58.2

        (Comeron et al., 2012)
        Left of (cM)
        Right of (cM)
        Notes
        Stocks and Reagents
        Stocks (7)
        Genomic Clones (22)
        cDNA Clones (348)
         

        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
        Other 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
           

          • New stable cell line derived from S1 : Stable S1 cell lines containing inducible transgenes for IntS6, defl (also known as IntS7), IntS12, asun (also known as IntS13), or IntS14 were created. They are denoted DL1 pMtnA IntS6, DL1 pMtnA IntS7, DL1 pMtnA IntS12, DL1 pMtnA IntS13, and DL1 pMtnA IntS14 by the author.

            (Fujiwara et al., 2023)
          Other Comments

          Mat89Bb is required for normal nucleus-centrosome coupling in primary spermatocytes and for the normal attachment between the nucleus and the basal body in postmeiotic spermatids. Mat89Bb is also required for the normal perinuclear localisation of dynein-dynactin complexes in both spermatocytes and spermatids.

          (Anderson et al., 2009)

          Putative substrate for the kinase png.

          (Lee et al., 2005)
          Relationship to Other Genes
          Source for database merge of
          Additional comments
          Nomenclature History
          Source for database identify of

          Source for identity of: asun Mat89Bb

          (Anderson et al., 2009)
          Nomenclature comments
          Etymology

          The gene is named "asunder" (set aprt in position or space) based on the mutant phenotype of a striking loss of association between asters and the nucleus during prophase in primary spermatocytes.

          (Anderson et al., 2009)
          Synonyms and Secondary IDs (12)
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 34 )
          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.
          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
          Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
          Flygut - An atlas of the Drosophila adult midgut
          FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
          iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
          MIST (protein-protein) - An integrated Molecular Interaction Database
          Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
          References (43)