General Information
Symbol
Dmel\Asx
Species
D. melanogaster
Name
Additional sex combs
Annotation Symbol
CG8787
Feature Type
FlyBase ID
FBgn0261823
Gene Model Status
Stock Availability
Gene Snapshot
Additional sex combs (Asx) is a chromatin binding protein with His2A deubiquitination activity. It genetically interacts with Pc and sxc and represses homeotic genes transcription. Asx contributes to antennal development and embryonic cell cycle. [Date last reviewed: 2017-06-15]
Also Known As
xen
Genomic Location
Cytogenetic map
Sequence location
2R:14,503,958..14,512,493 [+]
Recombination map
2-71
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
GO Summary Ribbons
Families, Domains and Molecular Function
Gene Group Membership (FlyBase)
Protein Family (UniProt, Sequence Similarities)
Belongs to the Asx family. (Q9V727)
Molecular Function (see GO section for details)
Summaries
Gene Group Membership
POLYCOMB REPRESSIVE DEUBIQUITINASE -
The Polycomb repressive deubiquitinase (PR-DUB) removes monoubiquitin from histone H2A. (Adapted from FBrf0210760).
UniProt Contributed Function Data
Atypical Polycomb group protein, which may be involved in both Polycomb group (PcG) and trithorax group (trxG) complexes. Non-catalytic component of the PR-DUB complex, a complex that specifically mediates deubiquitination of histone H2A monoubiquitinated at 'Lys-118' (H2AK118ub1). Does not deubiquitinate monoubiquitinated histone H2B. Required to maintain the transcriptionally repressive state of homeotic genes throughout development. The PR-DUB complex has weak or no activity toward 'Lys-48'- and 'Lys-63'-linked polyubiquitin chains. PcG and trxG proteins act by forming multiprotein complexes, which are respectively required to maintain the transcriptionally repressive and transcriptionally active state of homeotic genes throughout development. PcG and trxG protein complexes are not required to initiate repression and activation, but to maintain it during later stages of development.
(UniProt, Q9V727)
Phenotypic Description from the Red Book (Lindsley and Zimm 1992)
Asx: Additional sex combs
Asx/+ males have extra sex-comb teeth on meso- and metathoracic legs (e.g., Asx6 -> 11.8 sex-comb teeth per prothoracic, 0.5 per mesothoracic, and 0.2 per metathoracic leg). Homozygote embryonic lethal. Abdominal denticles in head and thoracic segments; abdominal segments 1-7 transformed into more posterior segments. With respect to the degree of transformation to more posterior structures, Df(2R)trix/Df(2R)trix > Df(2R)trix/Asx5 > Asx5/Asx5, indicating that Asx5 is hypomorphic mutation. Similar genotypes derived from Df(2R)trix/Asx5 oocytes achieved by means of pole-cell transplantatiton display more severe transformation than their counterparts above derived from mothers carrying one dose of Asx+ (Breen and Duncan). In double mutant combinations with Pcl, Psc, or Scm shows strong posterior transformation of all segments and failure of head involution. The presence of BXC+ required for expression of phenotype.
Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
4

Please see the GBrowse view of Dmel\Asx or the JBrowse view of Dmel\Asx 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
Comments on Gene Model
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.46
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0087493
6643
1669
FBtr0100154
5062
1560
FBtr0332929
6377
1670
FBtr0332930
6809
1538
Additional Transcript Data and Comments
Reported size (kB)
7.0 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0086622
179.8
1669
6.30
FBpp0099505
168.7
1560
6.39
FBpp0305149
179.9
1670
6.30
FBpp0305150
166.2
1538
6.39
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)
1668 (aa); 182 (kD predicted)
Comments
External Data
Subunit Structure (UniProtKB)
Component of the PR-DUB complex, at least composed of calypso and Asx (PubMed:20436459). Interacts with tant (PubMed:11397012). Interacts with cyclin CycG (PubMed:25995770).
(UniProt, Q9V727)
Domain
Contains two Leu-Xaa-Xaa-Leu-Leu (LXXLL) motifs, which may be required for an association with nuclear receptors.
(UniProt, Q9V727)
Crossreferences
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\Asx using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (19 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000339786
(assigned by GO_Central )
inferred from electronic annotation with InterPro:IPR026905
(assigned by InterPro )
Biological Process (12 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000339786
(assigned by GO_Central )
non-traceable author statement
(assigned by UniProt )
inferred from biological aspect of ancestor with PANTHER:PTN000339786
(assigned by GO_Central )
non-traceable author statement
(assigned by UniProt )
traceable author statement
Cellular Component (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
inferred from physical interaction with FLYBASE:calypso; FB:FBgn0262166
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
colocalizes_with nuclear chromatin
inferred from biological aspect of ancestor with PANTHER:PTN000339786
(assigned by GO_Central )
colocalizes_with nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000339786
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000339786
(assigned by GO_Central )
Expression Data
Transcript Expression
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Asx transcript is present at high levels in 0-1.5 hr embryos. Levels decline at 1.5-3 hours, then rise again. Levels are low in larvae, and increase in pupae and adults. Three transcripts are detected in adult males. Asx transcript is abundant in nurse cells, but absent from stage S10 oocytes. In the blastoderm stage embryo, expression is detected in a broad band in the anterior, and a narrower band in the posterior of the embryo. The transcript is ubiquitous during the rest of embryogenesis, though higher levels are detected in the neurectoderm and later in the central nervous system.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Asx protein is first detected in the cellular blastoderm embryo, in both the nucleus and cytoplasm. The protein may be at slightly higher levels in the anterior of the embryo. Later in embryogenesis, the protein is ubiquitous but non-uniform, with higher levels detected in the neurectoderm, and later in the central nervous system. Asx protein localizes to about 90 sites in polytene chromosomes.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
inferred from physical interaction with FLYBASE:calypso; FB:FBgn0262166
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Asx in GBrowse 2
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
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Images
Alleles, Insertions, Transgenic Constructs and Phenotypes
Classical and Insertion Alleles ( 41 )
For All Classical and Insertion Alleles Show
 
Allele of Asx
Class
Mutagen
Associated Insertion
Stocks
Known lesion
    0
    --
    Other relevant insertions
    miscellaneous insertions
    Name
    Expression Data
    Transgenic Constructs ( 5 )
    Deletions and Duplications ( 13 )
    Summary of Phenotypes
    For more details about a specific phenotype click on the relevant allele symbol.
    Lethality
    Allele
    Other Phenotypes
    Allele
    Phenotype manifest in
    Allele
    denticle belt & thorax
    Orthologs
    Human Orthologs (via DIOPT v7.1)
    Homo sapiens (Human) (3)
    Species\Gene Symbol
    Score
    Best Score
    Best Reverse Score
    Alignment
    Complementation?
    Transgene?
    9 of 15
    Yes
    Yes
    9 of 15
    Yes
    Yes
    9 of 15
    Yes
    Yes
    Model Organism Orthologs (via DIOPT v7.1)
    Mus musculus (laboratory mouse) (3)
    Species\Gene Symbol
    Score
    Best Score
    Best Reverse Score
    Alignment
    Complementation?
    Transgene?
    9 of 15
    Yes
    Yes
    8 of 15
    No
    Yes
    7 of 15
    No
    Yes
    Rattus norvegicus (Norway rat) (2)
    4 of 13
    Yes
    Yes
    1 of 13
    No
    Yes
    Xenopus tropicalis (Western clawed frog) (3)
    4 of 12
    Yes
    Yes
    3 of 12
    No
    Yes
    1 of 12
    No
    Yes
    Danio rerio (Zebrafish) (2)
    9 of 15
    Yes
    Yes
    8 of 15
    No
    Yes
    Caenorhabditis elegans (Nematode, roundworm) (0)
    No orthologs reported.
    Arabidopsis thaliana (thale-cress) (0)
    No orthologs reported.
    Saccharomyces cerevisiae (Brewer's yeast) (0)
    No orthologs reported.
    Schizosaccharomyces pombe (Fission yeast) (0)
    No orthologs reported.
    Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091901T0 )
    Organism
    Common Name
    Gene
    AAA Syntenic Ortholog
    Multiple Dmel Genes in this Orthologous Group
    Drosophila melanogaster
    fruit fly
    Drosophila suzukii
    Spotted wing Drosophila
    Drosophila simulans
    Drosophila sechellia
    Drosophila erecta
    Drosophila yakuba
    Drosophila ananassae
    Drosophila pseudoobscura pseudoobscura
    Drosophila persimilis
    Drosophila willistoni
    Drosophila virilis
    Drosophila mojavensis
    Drosophila grimshawi
    Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG0915010N )
    Organism
    Common Name
    Gene
    Multiple Dmel Genes in this Orthologous Group
    Musca domestica
    House fly
    Glossina morsitans
    Tsetse fly
    Lucilia cuprina
    Australian sheep blowfly
    Lucilia cuprina
    Australian sheep blowfly
    Aedes aegypti
    Yellow fever mosquito
    Culex quinquefasciatus
    Southern house mosquito
    Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W07ZC )
    Organism
    Common Name
    Gene
    Multiple Dmel Genes in this Orthologous Group
    Bombyx mori
    Silkmoth
    Danaus plexippus
    Monarch butterfly
    Heliconius melpomene
    Postman butterfly
    Apis florea
    Little honeybee
    Apis mellifera
    Western honey bee
    Bombus impatiens
    Common eastern bumble bee
    Bombus terrestris
    Buff-tailed bumblebee
    Linepithema humile
    Argentine ant
    Megachile rotundata
    Alfalfa leafcutting bee
    Nasonia vitripennis
    Parasitic wasp
    Nasonia vitripennis
    Parasitic wasp
    Dendroctonus ponderosae
    Mountain pine beetle
    Tribolium castaneum
    Red flour beetle
    Pediculus humanus
    Human body louse
    Rhodnius prolixus
    Kissing bug
    Cimex lectularius
    Bed bug
    Acyrthosiphon pisum
    Pea aphid
    Zootermopsis nevadensis
    Nevada dampwood termite
    Zootermopsis nevadensis
    Nevada dampwood termite
    Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X07VA )
    Organism
    Common Name
    Gene
    Multiple Dmel Genes in this Orthologous Group
    Strigamia maritima
    European centipede
    Ixodes scapularis
    Black-legged tick
    Stegodyphus mimosarum
    African social velvet spider
    Stegodyphus mimosarum
    African social velvet spider
    Stegodyphus mimosarum
    African social velvet spider
    Tetranychus urticae
    Two-spotted spider mite
    Daphnia pulex
    Water flea
    Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0C66 )
    Organism
    Common Name
    Gene
    Multiple Dmel Genes in this Orthologous Group
    Strongylocentrotus purpuratus
    Purple sea urchin
    Ciona intestinalis
    Vase tunicate
    Ciona intestinalis
    Vase tunicate
    Ciona intestinalis
    Vase tunicate
    Gallus gallus
    Domestic chicken
    Gallus gallus
    Domestic chicken
    Gallus gallus
    Domestic chicken
    Human Disease Model Data
    FlyBase Human Disease Model Reports
      Alleles Reported to Model Human Disease (Disease Ontology)
      Download
      Models ( 0 )
      Allele
      Disease
      Evidence
      References
      Interactions ( 0 )
      Allele
      Disease
      Interaction
      References
      Comments ( 0 )
       
      Human Orthologs (via DIOPT v7.1)
      Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
      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
      esyN Network Diagram
      esyN Network Key:
      Suppression
      Enhancement

      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
      External Data
      Subunit Structure (UniProtKB)
      Component of the PR-DUB complex, at least composed of calypso and Asx (PubMed:20436459). Interacts with tant (PubMed:11397012). Interacts with cyclin CycG (PubMed:25995770).
      (UniProt, Q9V727 )
      Linkouts
      DroID - A comprehensive database of gene and protein interactions.
      InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
      Pathways
      Gene Group - Pathway Membership (FlyBase)
      External Data
      Linkouts
      Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
      Genomic Location and Detailed Mapping Data
      Chromosome (arm)
      2R
      Recombination map
      2-71
      Cytogenetic map
      Sequence location
      2R:14,503,958..14,512,493 [+]
      FlyBase Computed Cytological Location
      Cytogenetic map
      Evidence for location
      51A5-51C1
      Left limit from inclusion within Df(2R)03072 (FBrf0093664) Right limit from inclusion within Df(2R)03072 (FBrf0093664)
      Experimentally Determined Cytological Location
      Cytogenetic map
      Notes
      References
      51A1-51B4
      (determined by in situ hybridisation)
      Experimentally Determined Recombination Data
      Left of (cM)
      Right of (cM)
      Notes
      Located on 2R.
      Stocks and Reagents
      Stocks (14)
      Genomic Clones (23)
      cDNA Clones (27)
       

      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 GBrowse for alignment of the cDNAs and ESTs to the gene model.

      cDNA clones, fully sequences
      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)
        RNAi and Array Information
        Linkouts
        DRSC - Results frm RNAi screens
        GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
        Antibody Information
        Laboratory Generated Antibodies
        Commercially Available Antibodies
         
        Other Information
        Relationship to Other Genes
        Source for database identify of
        Source for identity of: Asx CG8787
        Source for database merge of
        Additional comments
        Other Comments
        Mutant analysis demonstrates functional interactions between Pc, sxc and Asx in vivo.
        Asx is required for both the activation and repression of homeotic loci.
        Asx binds to multiple sites on polytene chromosomes, 70% of which overlap those of Pc, ph-p, ph-d and Pcl, 30% of which are unique.
        Asx is required in the central nervous system to regulate Ubx.
        Identification: Genetic screen for autosomal mutations that produce blisters in somatic wing clones. 1 allele of xen has been isolated.
        In an effort to subdivide the Pc-group genes functionally, the phenotypes of adult flies heterozygous for every pairwise combination of Pc-group mutation were examined. Asx, Pc, Pcl, Psc, Scm and Sce have similar functions in some imaginal tissues. Genetic interactions have been demonstrated between esc, Asx, E(Pc), Pcl, E(z) and sxc. Most duplications of Pc-group genes neither exhibit anterior transformations nor suppress the extra sex comb phenotype of Pc-group mutations, suggesting that not all Pc-group genes behave as predicted by the mass action model.
        Sections of the Scr regulatory region may be important for regulation of Scr by Polycomb- and trithorax-group genes.
        Mosaic and expression pattern analysis reveals that the Pc-group genes do not act only in a common complex or pathway: they must have some independent functions.
        The bithorax complex genes are regulated by the Pc-group of genes, acting via 'Pc-group response elements' (PREs), that can work even when removed from the normal bithorax complex context. The Pc-group products apparently provide stable memory or imprinting of boundaries which are specified by gap and pair-rule regulators.
        Mutations of genes in the Polycomb group (esc, E(z), Pc, ph-p, ph-d, Scm, Pcl, Sce, Asx, Psc, pho and Antp) cause abnormal segmental development due to the ectopic expression of abd-A and Abd-B. Embryos lacking both maternal and zygotic Asx product were generated to determine abd-A and Abd-B expression patterns.
        Embryonic and adult phenotypes suggest that Asx is required zygotically for determination of segment number and polarity. A construct carrying an eve promoter fused to a Ecol\lacZ gene was used to demonstrate that 40% of the mutant Asx embryos exhibit some ectopic expression of eve. This suggests Asx is required for the normal expression of eve.
        The Pc-group genes are negative regulators of homeotic genes.
        Pole cell transplantation techniques demonstrate that Asx is maternally expressed and is required for normal the bithorax complex expression during embryogenesis.
        A member of the Polycomb group of genes. Asx function is required for the regulation of the bithorax complex during embryonic development. In its absence abd-A and Abd-B are ectopically expressed, leading to the conclusion that Asx is a negative regulator of the bithorax complex.
        Asx/+ males have extra sex-comb teeth on meso- and metathoracic legs. Homozygote embryonic lethal. Abdominal denticles in head and thoracic segments; abdominal segments 1-7 transformed into more posterior segments. In double mutant combinations with Pcl, Psc, or Scm shows strong posterior transformation of all segments and failure of head involution. The presence of the bithorax complex+ required for expression of phenotype.
        Origin and Etymology
        Discoverer
        Jurgens.
        Etymology
        Identification
        External Crossreferences and Linkouts ( 47 )
        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/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
        UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
        Other crossreferences
        Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
        Linkouts
        Drosophila Genomics Resource Center - Drosophila Genomics Resource Center 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.
        Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
        FlyMine - An integrated database for Drosophila genomics
        GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
        iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
        InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
        KEGG Genes - Molecular building blocks of life in the genomic space.
        modMine - A data warehouse for the modENCODE project
        Synonyms and Secondary IDs (11)
        Reported As
        Secondary FlyBase IDs
        • FBgn0000142
        • FBgn0020770
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        References (163)