FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\rux
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General Information
Symbol
Dmel\rux
Species
D. melanogaster
Name
roughex
Annotation Symbol
CG4336
Feature Type
protein_coding_gene
FlyBase ID
FBgn0003302
Gene Model Status
Current
Stock Availability
19 publicly available
Gene Summary
Negative regulator of both mitosis and meiosis. Required for the establishment of the G1 phase in the developing eye. (UniProt, P50445)
Contribute a Gene Snapshot for this gene.
All Summaries
Key Links
Genomic Location
Cytogenetic map
5D2-5D2
Sequence location
Recombination map
1-15
RefSeq locus
NC_004354 REGION:6035753..6037800
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 (4 terms)
Molecular Function (0 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
involved_in mitotic cell cycle
inferred from mutant phenotype
(Foley and Sprenger, 2001)
involved_in regulation of cell cycle
inferred from mutant phenotype
(Avedisov et al., 2000)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
involved_in regulation of exit from mitosis
traceable author statement
(Lee and Orr-Weaver, 2003)
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
located_in nucleus
inferred from direct assay
(Avedisov et al., 2000, Foley et al., 1999)
Terms Based on Predictions or Assertions (0 terms)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Protein Function (UniProtKB)
Negative regulator of both mitosis and meiosis. Required for the establishment of the G1 phase in the developing eye.
(UniProt, P50445)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
rux: roughex
Eyes smaller than wild type and uniformly rough. Male sterile. RK2.
Summary (Interactive Fly)

novel regulator of cell cycle - a cyclin dependent kinase inhibitor that specifically inhibits Cdk1-CycA - required to inhibit Cdk1-CycA kinase activity during metaphase, thereby facilitating the transition to anaphase

Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\rux 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 P50445)

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 5.46

Low-frequency RNA-Seq exon junction(s) not annotated.

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
rux-RA
FBtr0070889
NM_078498
1420
335
rux-RB
FBtr0307288
NM_001258611
1319
335
Additional Transcript Data and Comments
Reported size (kB)

1.9, 1.3 (northern blot)

(Thomas et al., 1994)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
rux-PA
FBpp0070854
37.9
335
5.13
P50445
NP_511053
AAF46113
rux-PB
FBpp0298289
37.9
335
5.13
P50445
NP_001245540
AFH07254
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

335 aa isoforms: rux-PA, rux-PB
Additional Polypeptide Data and Comments
Reported size (kDa)

335 (aa)

(Thomas et al., 1994)
Comments
External Data
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\rux 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).

4.62

Transcript Expression
Polypeptide Expression
distribution deduced from reporter or direct label
Stage
Tissue/Position (including subcellular localization)
Reference
spermatogenesis & adult stage | male
testis
(Franklin-Dumont et al., 2007)
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in nucleus
inferred from direct assay
(Avedisov et al., 2000, Foley et al., 1999)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\rux 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
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
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 9-10
  • Stages(s) 11-12
  • Stages(s) 13-16
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 13 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 17 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of rux
Transgenic constructs containing regulatory region of rux
Aberrations (Deficiencies and Duplications) ( 20 )
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) (0)
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) (0)
Mus musculus (laboratory mouse) (0)
Xenopus tropicalis (Western clawed frog) (0)
Danio rerio (Zebrafish) (0)
Caenorhabditis elegans (Nematode, roundworm) (0)
Anopheles gambiae (African malaria mosquito) (0)
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:rux. 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?
    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
    rux
    enhanceable
    ast
    (Thomas et al., 1994)
    rux
    suppressible
    CycA
    (Dong et al., 1997, Gonczy et al., 1994, Thomas et al., 1994)
    rux
    suppressible
    CycB
    (Gonczy et al., 1994)
    rux
    enhanceable
    ebi
    (Dong et al., 1999)
    rux
    enhanceable
    Ras85D
    (Thomas et al., 1994)
    rux
    suppressible
    Rca1
    (Dong et al., 1997)
    rux
    suppressible
    S(rux)2C
    (Dong et al., 1997)
    rux
    enhanceable
    S
    (Thomas et al., 1994)
    rux
    suppressible
    stg
    (Dong et al., 1997, Thomas et al., 1994)
    rux
    suppressible
    swm
    (Casso et al., 2008, Dong et al., 1997)
    rux
    suppressible
    twe
    (Gonczy et al., 1994, Thomas et al., 1994)
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    CycA
    enhanceable
    rux
    (Foley and Sprenger, 2001)
    CycA
    suppressible
    rux
    (Foley and Sprenger, 2001)
    CycA
    suppressible
    rux|Cdk1
    (Foley et al., 1999)
    CycA|Cdk1
    suppressible
    rux
    (Foley et al., 1999)
    CycE
    enhanceable
    rux
    (Secombe et al., 1998)
    CycE
    suppressible
    rux
    (Brumby et al., 2004, Secombe et al., 1998)
    enc
    suppressible
    rux
    (Ohlmeyer and Schupbach, 2003)
    Myt1
    suppressible
    rux
    (Willms et al., 2020, Varadarajan et al., 2016)
    upd1
    suppressible
    rux
    (Mukherjee et al., 2006)
    Wee1
    enhanceable
    rux
    (Price et al., 2002)
    External Data
    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
    1-15
    Cytogenetic map
    5D2-5D2
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    5D2-5D2
    Limits computationally determined from genome sequence between P{EP}EP1457EP1457&P{EP}CG5928EP1083 and P{EP}EP1311&P{EP}EP1384
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    Experimentally Determined Recombination Data
    Location

    1-15

    (FlyBase, 2016)

    1-15.8

    (Comeron et al., 2012)

    1-15.0

    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (19)
    Genomic Clones (16)
     

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

    cDNA Clones (23)
     

    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
       
        Other Comments

        Shows particularly robust cycling of transcription in adult heads, as assessed by expression analysis using high density oligonucleotide arrays with probe generated during three 12-point time course experiments over the course of 6 days.

        (Claridge-Chang et al., 2001)

        rux is involved in mitotic exit.

        (Foley and Sprenger, 2001)

        rux is a distinct and specialised cell cycle inhibitor for CycA dependant kinase activity.

        (Avedisov et al., 2000)

        rux is a cyclin dependent kinase inhibitor (CKI) specific for mitotic cyclin-cdk complexes.

        (Foley et al., 1999)

        The mechanism by which dpp mediates G1 arrest in the morphogenetic furrow does not require rux.

        (Horsfield et al., 1998)

        In vitro studies demonstrate rux binds CycE but does not inhibit CycE-Cdk activity directly. Overexpression studies suggest the ectopic rux converts the mitotic cycle into an endoreplication cycle by down-regulating CycA-associated kinase activity.

        (Thomas et al., 1997)

        rux acts genetically to negatively regulate CycA and stg.

        (Thomas et al., 1995)

        rux is a dose-dependent regulator of meiosis II during spermatogenesis. rux appears to act through CycA during premeiotic G2 to regulate meiosis II.

        (Gonczy et al., 1994)

        rux functions as a negative regulator of G1 progression in the developing eye.

        (Thomas et al., 1994)

        The rux mutant phenotype is due to a developmental defect: pattern formation defect.

        (Renfranz and Benzer, 1989)
        Relationship to Other Genes
        Source for database merge of
        Additional comments
        Nomenclature History
        Source for database identify of

        Source for identity of: rux CG4336

        (Zhao, 1999.11)
        Nomenclature comments
        Etymology
        Synonyms and Secondary IDs (6)
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 38 )
        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
        UniProt/TrEMBL - Automatically annotated and unreviewed 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
        Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
        Flygut - An atlas of the Drosophila adult midgut
        Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
        MIST (genetic) - An integrated Molecular Interaction Database
        MIST (protein-protein) - An integrated Molecular Interaction Database
        References (129)