FB2025_02 , released April 17, 2025
Gene: Dmel\Cse1
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General Information
Symbol
Dmel\Cse1
Species
D. melanogaster
Name
Chromosome segregation 1
Annotation Symbol
CG13281
Feature Type
protein_coding_gene
FlyBase ID
FBgn0022213
Gene Model Status
Current
Stock Availability
13 publicly available
Gene Summary
Export receptor for importin alpha. Mediates importin-alpha re-export from the nucleus to the cytoplasm after import substrates have been released into the nucleoplasm (By similarity). (UniProt, Q9XZU1)
Contribute a Gene Snapshot for this gene.
All Summaries
Also Known As

Cas, Dcas, l(2)k03902, CAS/CSE1 segregation protein

Key Links
Genomic Location
Cytogenetic map
36B1-36B2
Sequence location
Recombination map
2-53
RefSeq locus
NT_033779 REGION:16821156..16825207
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 (8 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables nuclear export signal receptor activity
inferred from biological aspect of ancestor with PANTHER:PTN000108304
(GO Reference Genome Project, 2011-)
enables small GTPase binding
inferred from electronic annotation with InterPro:IPR001494, InterPro:IPR005043
(InterPro Project Members, 2004-)
Biological Process (3 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
involved_in protein export from nucleus
inferred from mutant phenotype
(Tekotte et al., 2002)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
involved_in intracellular protein transport
inferred from electronic annotation with InterPro:IPR001494, InterPro:IPR013713
(InterPro Project Members, 2004-)
involved_in protein export from nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000108304
(GO Reference Genome Project, 2011-)
involved_in protein import into nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000108134
(GO Reference Genome Project, 2011-)
Cellular Component (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
located_in cytosol
inferred from high throughput direct assay
(Lye et al., 2014)
located_in nuclear envelope
inferred from high throughput direct assay
(Lye et al., 2014)
located_in nucleoplasm
inferred from high throughput direct assay
(Lye et al., 2014)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in cytosol
inferred from biological aspect of ancestor with PANTHER:PTN000108134
(GO Reference Genome Project, 2011-)
is_active_in nuclear envelope
inferred from biological aspect of ancestor with PANTHER:PTN000108134
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the XPO2/CSE1 family. (Q9XZU1)
Protein Signatures (InterPro)
Summaries
Gene Group (FlyBase)
BETA IMPORTINS -
Importin-β family proteins are nucleocytoplasmic transport receptors that import and export proteins and RNAs through the nuclear pores. In some import/export pathways, importin-β directly interacts with the cargo. For cargo containing a classical nuclear localization (cNLS) signal, importin-α acts as an adaptor linking the cargo to importin-β. (Adapted from FBrf0235243, FBrf0131350 and PMID:21029754).
Protein Function (UniProtKB)
Export receptor for importin alpha. Mediates importin-alpha re-export from the nucleus to the cytoplasm after import substrates have been released into the nucleoplasm (By similarity).
(UniProt, Q9XZU1)
Summary (Interactive Fly)

signaling protein - functions in axonal fasciculation and pathfinding - functions with integrins to specify axon guidance - functions in nuclear transport in the Notch pathway

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

Please see the JBrowse view of Dmel\Cse1 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 Q9XZU1)

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 5.45

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Cse1-RA
FBtr0080930
NM_078864
3237
975
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
Cse1-PA
FBpp0080484
110.1
975
5.50
Q9XZU1
NP_523588
AAF53575
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)

Binds with high affinity to importin-alpha only in the presence of RanGTP.

(UniProt, Q9XZU1)
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\Cse1 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).

-0.29

Transcript Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult head
(Aradska et al., 2015)
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytosol
inferred from high throughput direct assay
(Lye et al., 2014)
located_in nuclear envelope
inferred from high throughput direct assay
(Lye et al., 2014)
located_in nucleoplasm
inferred from high throughput direct assay
(Lye et al., 2014)
Expression Deduced from Reporters
Reporter: PBac{602.P.SVS-1}Cse1CPTI000199
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult brain cell body rind

Comment: low expression in the adult brain cortex

(Knowles-Barley, 2011.8.24)
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Cse1 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 ( 18 )
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 Cse1
Transgenic constructs containing regulatory region of Cse1
Aberrations (Deficiencies and Duplications) ( 4 )
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) (5)
Hsap\CSE1L
14 of 14
Yes
Yes
Hsap\IPO9
2 of 14
No
No
Hsap\IPO11
2 of 14
No
No
Hsap\IPO7
1 of 14
No
No
Hsap\IPO8
1 of 14
No
No
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) (5)
Rnor\Cse1l
14 of 14
Yes
Yes
Rnor\Ipo9
2 of 14
No
No
Rnor\Ipo7
1 of 14
No
No
Rnor\Ipo8
1 of 14
No
No
Rnor\Ipo11
1 of 14
No
No
Mus musculus (laboratory mouse) (5)
Mmus\Cse1l
13 of 14
Yes
Yes
Mmus\Ipo9
2 of 14
No
No
Mmus\Ipo11
2 of 14
No
No
Mmus\Ipo7
1 of 14
No
No
Mmus\Ipo8
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (6)
Xtro\cse1l
11 of 13
Yes
Yes
Xtro\bdh1a
9 of 13
No
Yes
Xtro\allc
1 of 13
No
Yes
Xtro\ipo8
1 of 13
No
No
Xtro\ipo9
1 of 13
No
No
Xtro\ipo11
1 of 13
No
No
Danio rerio (Zebrafish) (4)
Drer\cse1l
14 of 14
Yes
Yes
Drer\ipo7
1 of 14
No
No
Drer\ipo8
1 of 14
No
No
Drer\ipo9
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (2)
Cele\xpo-2
13 of 14
Yes
Yes
Cele\imb-2
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (4)
Agam\AgaP_AGAP010711
12 of 12
Yes
Yes
Agam\AgaP_AGAP011156
2 of 12
No
No
Agam\AgaP_AGAP002970
1 of 12
No
No
Agam\AgaP_AGAP007299
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (6)
Atha\AT2G46520
13 of 13
Yes
Yes
Atha\AT1G26170
2 of 13
No
No
Atha\AT3G08960
1 of 13
No
No
Atha\AT3G17340
1 of 13
No
No
Atha\AT3G59020
1 of 13
No
No
Atha\SAD2
1 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (5)
Scer\CSE1
13 of 13
Yes
Yes
Scer\KAP114
2 of 13
No
No
Scer\KAP120
2 of 13
No
No
Scer\NMD5
1 of 13
No
No
Scer\SXM1
1 of 13
No
No
Schizosaccharomyces pombe (Fission yeast) (4)
Spom\kap109
12 of 12
Yes
Yes
Spom\kap113
2 of 12
No
No
Spom\kap114
2 of 12
No
No
Spom\SPCC550.11
1 of 12
No
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Cse1. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (3)
Ipo9
4 of 13
Impβ11
3 of 13
msk
3 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 ( 2 )
    Allele
    Disease
    Interaction
    References
    Cse1k03902
    ameliorates  autosomal dominant cerebellar ataxia
    modeled by Hsap\ATXN8OSCTG112.UAS
    (Mutsuddi et al., 2004)
    Cse1JF02972
    exacerbates  cancer
    modeled by RbfHMS03004
    (Parkhitko et al., 2021)
    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?
    CSE1L; chromosome segregation 1 like
    14 of 14
    601342
        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
        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
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        Kap-α1
        enhanceable
        Cse1
        (Ratan et al., 2008)
        Kap-α1
        suppressible
        Cse1
        (Ratan et al., 2008)
        External Data
        Subunit Structure (UniProtKB)
        Binds with high affinity to importin-alpha only in the presence of RanGTP.
        (UniProt, Q9XZU1 )
        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
        KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
        Class of Gene
        Genomic Location and Detailed Mapping Data
        Chromosome (arm)
        2L
        Recombination map
        2-53
        Cytogenetic map
        36B1-36B2
        Sequence location
        FlyBase Computed Cytological Location
        Cytogenetic map
        Evidence for location
        36B1-36B2
        Limits computationally determined from genome sequence between P{lacW}Mhck10423&P{lacW}Cask03902 and P{lacW}Aac11k06710
        Experimentally Determined Cytological Location
        Cytogenetic map
        Notes
        References
        36B1-36B2
        (Tekotte, 1999.5.6, BDGP Project Members, 1994-1999)
        36B1-36B2
        (determined by in situ hybridisation)
        (Spradling et al., 1999)
        Experimentally Determined Recombination Data
        Location

        2-53

        (FlyBase, 2016)

        2-51.3

        (Comeron et al., 2012)
        Left of (cM)
        Right of (cM)
        Notes
        Stocks and Reagents
        Stocks (13)
        Genomic Clones (15)
         

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

        cDNA Clones (39)
         

        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

              S2 cells treated with dsRNA generated against this gene show reduced phagocytosis of Candida albicans compared to untreated cells.

              (Stroschein-Stevenson et al., 2006)

              Cas is a ubiquitous export receptor that seems to be required in all tissues.

              (Tekotte et al., 2002)
              Relationship to Other Genes
              Source for database merge of
              Additional comments
              Nomenclature History
              Source for database identify of

              Source for identity of: Cse1 Cas

              (FlyBase, 2009-)
              Nomenclature comments

              Renamed from 'Cas' to 'Cse1' in order to minimise confusion with the unrelated 'cas' gene (which only differed by case) and to be consistent with the nomenclature used in other organisms for orthologs of this gene.

              (FlyBase, 2009-)
              Etymology
              Synonyms and Secondary IDs (13)
              Datasets (0)
              Study focus (0)
              Experimental Role
              Project
              Project Type
              Title
              Study result (0)
              Result
              Result Type
              Title
              External Crossreferences and Linkouts ( 41 )
              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
              iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
              Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
              KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
              MIST (genetic) - An integrated Molecular Interaction Database
              MIST (protein-protein) - An integrated Molecular Interaction Database
              References (85)