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General Information
Symbol
Dmel\Pen
Species
D. melanogaster
Name
Pendulin
Annotation Symbol
CG4799
Feature Type
FlyBase ID
FBgn0287720
Gene Model Status
Stock Availability
Gene Snapshot
Pendulin (Pen) encodes a member of the Importin-alpha protein family, an essential adaptor in the Ran-GTP nuclear transport cycle binding a cargo protein to the nuclear import receptor encoded by Fs(2)Ket. In addition to nuclear protein import, the product of Pen is involved in centrosome duplication, mitotic spindle dynamics, nuclear envelope assembly, ring canal formation in the female germline, geotaxic behaviour and perception of pain. [Date last reviewed: 2019-03-14]
Also Known As

imp-α2, importin α2, mushroom body miniature B, mbmB, oho31

Key Links
Genomic Location
Cytogenetic map
Sequence location
2L:10,056,900..10,060,095 [+]
Recombination map

2-39

RefSeq locus
NT_033779 REGION:10056900..10060095
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (13 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR002652, InterPro:IPR024931
(assigned by InterPro )
inferred from biological aspect of ancestor with PANTHER:PTN000593662
(assigned by GO_Central )
Biological Process (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000593662
(assigned by GO_Central )
Cellular Component (5 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000593662
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000593662
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000593662
(assigned by GO_Central )
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the importin alpha family. (P52295)
Summaries
Gene Group (FlyBase)
ALPHA IMPORTINS -
The importin-α proteins mediate nuclear import by acting as adaptors that link classical nuclear localization signal containing proteins to importin-β. (Adapted from FBrf0149010, FBrf0207686 and PMID:15350979).
Protein Function (UniProtKB)
Binds specifically and directly to substrates containing either a simple or bipartite NLS motif. Promotes docking of import substrates to the nuclear envelope. Seems to act as a cytosolic receptor for both simple and bipartite NLS motifs (By similarity). It is required for normal cell proliferation and may serve as an adapter molecule to form complexes with other proteins. May act as a tumor suppressor in hematopoietic cells. May play a role in the nuclear import of karyophilic proteins and some of these may be required for the normal transmission and function of proliferative signals in the cells.
(UniProt, P52295)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
mbmB: mushroom-body-miniature-B (J.C. Hall)
Phenotype similar to that of mbm, but less extreme, and mushroom body defect occurs in both males and females; learning is weak in tests using olfactory stimuli.
Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\Pen 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

Gene model reviewed during 5.52

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0079937
2577
522
Additional Transcript Data and Comments
Reported size (kB)

2.8 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0079527
57.8
522
4.99
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)

522 (aa); 54 (kD observed); 57.8 (kD predicted)

522 (aa); 58 (kD predicted)

Comments
External Data
Subunit Structure (UniProtKB)

Forms a complex with importin beta subunit.

(UniProt, P52295)
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
Mapped Features

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

External Data
Crossreferences
Linkouts
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism | ubiquitous

Comment: reference states 0-2 hr AEL

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

Pen transcript is detected at high levels in early embryos (0-4 hr), and is barely detectable in late embryos and first instar larvae. Expression levels increase significantly in second larval instar, and is high in adults, particularly females.

The Pen transcript is detected at high levels in 0-2 hr embryos, decreases through gastrulation, and increases once more during germ band extension before disappearing in late embryogenesis. Expression is detected once more in late third instar larvae and persists through pupal and adult stages. Expression is particularly high in adult females. In situ localization experiments show the early embryonic Pen expression to be ubiquitous. By nuclear cycle 10-13, expression is most intense in the pole cells, but seven weak stripes of Pen expression are visible in the anterioposterior axis. At gastrulation, the stripes are stronger. At germ band extension, Pen transcripts are restricted to neuroblasts and ventral ectoderm. After germ band extension, Pen transcript is found in the ventral nerve cord, and the proliferating regions of the brain lobe. In third larval instar, staining is detected in the imaginal discs, and more weakly in the ring and lymph glands. Expression is also detectable in the brain hemispheres and the ventral ganglion.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

The location of Pen protein changes during the cell cycle. During interphase, Pen protein is found in the embryonic cytoplasm. At the G2-M transition, Pen protein translocates to the nucleus. A similar cell-cyle dependent redistribution is seen in larval CNS neuroblasts.

The localization of Pen protein changes during the cell cycle. The protein is cytoplasmic during interphase, and is nuclear at the onset of mitosis. During its nuclear localization, Pen protein does not appear to actively associate with chromatin.

Although the size of the in-vitro translated Pen protein agrees with the predicted size, two size classes are seen in immunoblots. Phosphatase treatment showed that phosphorylation was responsible for the differences in migration. In immunoblots, the faster migrating species was seen in all developmental stages. High levels were detected in ovaries and early embryos. Only low levels were detected during larval development. The slower migrating species was only detected in ovaries and preblastoderm embyros.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Pen 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
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 28 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 28 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Pen
Transgenic constructs containing regulatory region of Pen
Deletions and Duplications ( 0 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (8)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
12 of 15
Yes
Yes
8 of 15
No
Yes
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
1 of 15
No
Yes
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (9)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
7 of 15
Yes
Yes
7 of 15
Yes
Yes
6 of 15
No
Yes
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
2 of 15
No
No
1 of 15
No
Yes
1 of 15
No
Yes
Rattus norvegicus (Norway rat) (10)
11 of 13
Yes
Yes
6 of 13
No
Yes
6 of 13
No
Yes
3 of 13
No
No
3 of 13
No
No
3 of 13
No
No
3 of 13
No
No
2 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (8)
10 of 12
Yes
Yes
7 of 12
No
Yes
3 of 12
No
No
3 of 12
No
No
3 of 12
No
No
3 of 12
No
No
1 of 12
No
No
1 of 12
No
Yes
Danio rerio (Zebrafish) (8)
8 of 15
Yes
Yes
7 of 15
No
Yes
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
2 of 15
No
No
1 of 15
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (3)
3 of 15
Yes
No
2 of 15
No
No
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (11)
2 of 9
Yes
Yes
2 of 9
Yes
Yes
2 of 9
Yes
No
2 of 9
Yes
No
2 of 9
Yes
No
2 of 9
Yes
No
2 of 9
Yes
No
2 of 9
Yes
No
2 of 9
Yes
No
2 of 9
Yes
No
1 of 9
No
Yes
Saccharomyces cerevisiae (Brewer's yeast) (1)
4 of 15
Yes
No
Schizosaccharomyces pombe (Fission yeast) (2)
2 of 12
Yes
No
2 of 12
Yes
No
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG0919064V )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
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) ( EOG09150516 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Glossina morsitans
Tsetse fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W04WH )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis florea
Little honeybee
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Apis mellifera
Western honey bee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus impatiens
Common eastern bumble bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Bombus terrestris
Buff-tailed bumblebee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Megachile rotundata
Alfalfa leafcutting bee
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Tribolium castaneum
Red flour beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Pediculus humanus
Human body louse
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Rhodnius prolixus
Kissing bug
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Cimex lectularius
Bed bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Zootermopsis nevadensis
Nevada dampwood termite
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X03XA )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
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
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G095Z )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
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
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (3)
5 of 10
5 of 10
3 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    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 v8.0 and OMIM)
    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
    esyN Network Diagram
    Show neighbor-neighbor interactions:
    Select Layout:
    Legend:
    Protein
    RNA
    Selected Interactor(s)
    Interactions Browser

    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
    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)
    Forms a complex with importin beta subunit.
    (UniProt, P52295 )
    Linkouts
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    External Data
    Linkouts
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2L
    Recombination map

    2-39

    Cytogenetic map
    Sequence location
    2L:10,056,900..10,060,095 [+]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    31A1-31A2
    Left limit from in situ hybridisation (FBrf0067338) Right limit from in situ hybridisation (FBrf0067338)
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    31A1-31A2
    (determined by in situ hybridisation)
    31A-31A
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Location
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (26)
    Genomic Clones (24)
    cDNA Clones (260)
     

    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 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)
        Other clones
        RNAi and Array Information
        Linkouts
        Antibody Information
        Laboratory Generated Antibodies
        Commercially Available Antibodies
         
        Other Information
        Relationship to Other Genes
        Source for database identify of
        Source for database merge of

        Source for merge of: Pen anon-WO0140519.258

        Source for merge of: Pen l(2)k06324

        Source for merge of: Pen mbmB

        Additional comments

        Source for merge of Pen anon-WO0140519.258 was sequence comparison ( date:051113 ).

        FlyBase curator comment: Designation as "oho-31" is in error, the gene referred to as "oho-31" in FBrf0137017 is actually "Pen" (FBgn0011823). Confusion arose because the phenotypes of the "l(2)k14401" insertion line (the larval lethality and overgrown hematopoietic tissue phenotypes, which are caused by mutation in oho31, FBgn0010332) are separable from the insertion (which is in Pen) (see FBrf0149010).

        FlyBase curator comment: Designation as "oho31" is in error, the gene referred to as "oho31" in FBrf0103636 is actually "Pen" (FBgn0011823). Confusion arose because the phenotypes of the "l(2)k14401" insertion line (the larval lethality and overgrown hematopoietic tissue phenotypes, which are caused by mutation in oho31, FBgn0010332) are separable from the insertion (which is in Pen) (see FBrf0149010).

        FlyBase curator comment: Designation as "OHO31" is in error, the gene referred to as "OHO31" in FBrf0099880 is actually "Pen" (FBgn0011823). Confusion arose because the phenotypes of the "l(2)k14401" insertion line (the larval lethality and overgrown hematopoietic tissue phenotypes, which are caused by mutation in oho31, FBgn0010332) are separable from the insertion (which is in Pen) (see FBrf0149010).

        FlyBase curator comment: Designation as "oho31" is in error, the gene referred to as "oho31" in FBrf0088012 is actually "Pen" (FBgn0011823). Confusion arose because the phenotypes of the "l(2)k14401" insertion line (the larval lethality and overgrown hematopoietic tissue phenotypes, which are caused by mutation in oho31, FBgn0010332) are separable from the insertion (which is in Pen) (see FBrf0149010).

        FlyBase curator comment: Designation as "Oho31" is in error, the gene referred to as "Oho31" in FBrf0080740 is actually "Pen" (FBgn0011823). Confusion arose because the phenotypes of the "l(2)k14401" insertion line (the larval lethality and overgrown hematopoietic tissue phenotypes, which are caused by mutation in oho31, FBgn0010332) are separable from the insertion (which is in Pen) (see FBrf0149010).

        FlyBase curator comment: "oho31" phenotype (overgrown hematopoietic tissues and larval lethality) in the "l(2)k14401" insertion line is stated in FBrf0082252 to be due to an effect on the "Pen" gene, however, FBrf0149010 states that the "oho31" mutant phenotype is caused by a second site mutation separable from the insertion in "Pen" in the "l(2)k14401" insertion line.

        FlyBase curator comment: Designation as "OHO31" is in error, the gene referred to as "OHO31" in FBrf0079531 is actually "Pen" (FBgn0011823). Confusion arose because the phenotypes of the "l(2)k14401" insertion line (the larval lethality and overgrown hematopoietic tissue phenotypes, which are caused by mutation in oho31, FBgn0010332) are separable from the insertion (which is in Pen) (see FBrf0149010).

        FlyBase curator comment: Designation as "oho31" is in error, the gene referred to as "oho31" in FBrf0083561 is actually "Pen" (FBgn0011823). Confusion arose because the phenotypes of the "l(2)k14401" insertion line (the larval lethality and overgrown hematopoietic tissue phenotypes, which are caused by mutation in oho31, FBgn0010332) are separable from the insertion (which is in Pen) (see FBrf0149010).

        FlyBase curator comment: "oho31" phenotype (overgrown hematopoietic tissues and larval lethality) in the "l(2)k14401" insertion line is stated in FBrf0082712 to be due to an effect on the "Pen" gene, however, FBrf0149010 states that the "oho31" mutant phenotype is caused by a second site mutation separable from the insertion in "Pen" in the "l(2)k14401" insertion line.

        FlyBase curator comment: Designation as "oho31" is in error, the gene referred to as "Oho31" in FBrf0121419 is actually "Pen" (FBgn0011823). Confusion arose because the phenotypes of the "l(2)k14401" insertion line (the larval lethality and overgrown hematopoietic tissue phenotypes, which are caused by mutation in oho31, FBgn0010332) are separable from the insertion (which is in Pen) (see FBrf0149010).

        Other Comments

        dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.

        Pen may play a critical role in kel function by regulating its deposition on ring canals during their assembly.

        Pen is a nuclear localisation signal (NLS)-binding protein that is involved in the intercellular transport of NLS-containing proteins from the nurse cells to the oocytes.

        Loss of function mutations result in female sterility; homozygous females lay eggs which are smaller than wild-type and have open anterior ends. The oocytes are smaller than normal in stage 8 egg chambers and have an uneven surface, indicating that the early phase of cytoplasm transport is defective.

        Pen gene product is a cell cycle-dependent component of the nucleus that may be required for critical nuclear reactions occurring during the onset of mitosis.

        The Pen product may act as a cytosolic factor in nuclear protein transport. The cell cycle-dependent accumulation of Pen product in the nucleus indicates that this protein may be required for critical nuclear reactions occurring at the onset of mitosis.

        A sequence comparison and numerical analysis of the RRM-containing (RNA recognition motif) proteins suggests that functionally related RRM-containing proteins have significant sequence similarities in their RRMs.

        Pen is localised in the cytoplasm during interphase, rapidly translocates into the nucleus at G2/M-phase transition and remains in the vicinity of the chromatin throughout the M-phase.

        Behavioural and anatomical studies demonstrate that central brain lesions can be interpreted behaviourally.

        Origin and Etymology
        Discoverer
        Etymology
        Identification
        External Crossreferences and Linkouts ( 46 )
        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 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
        Synonyms and Secondary IDs (55)
        Reported As
        Symbol Synonym
        anon-WO0140519.258
        importin α2/pendulin
        l(2)k06324
        l(2)k14401
        mbmBN806
        rem
        Secondary FlyBase IDs
        • FBgn0267727
        • FBgn0011823
        • FBgn0014274
        • FBgn0026803
        • FBgn0044610
        • FBgn0022138
        • FBgn0002688
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        References (164)