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General Information
Symbol
Dmel\caup
Species
D. melanogaster
Name
caupolican
Annotation Symbol
CG10605
Feature Type
FlyBase ID
FBgn0015919
Gene Model Status
Stock Availability
Gene Snapshot
caupolican (caup) encodes a homeodomain-containing protein of the TALE subfamily and one of the three components of the Iroquois gene complex. caup product acts as a transcriptional regulator to control territorial and cell fate specification, cell sorting behaviour and pattern formation. It also plays a transcription independent cell-autonomous role in cell proliferation control. [Date last reviewed: 2018-09-20]
Also Known As
iro, iroquois, Iro-C
Key Links
Genomic Location
Cytogenetic map
Sequence location
3L:12,609,631..12,621,798 [+]
Recombination map
3-38
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
Belongs to the TALE/IRO homeobox family. (P54269)
Summaries
Gene Group (FlyBase)
TALE HOMEOBOX TRANSCRIPTION FACTORS -
TALE homeobox transcription factors are sequence-specific DNA binding proteins that regulate transcription. These proteins are characterized by an atypical homeodomain that possesses a three amino acid insertion in the first loop. (Adapted from FBrf0232555 and PMID:18797923).
Protein Function (UniProtKB)
Controls proneural and vein forming genes. Positive transcriptional controler of ac-sc (achaete-scute). May act as an activator that interacts with the transcriptional complex assembled on the ac and sc promoters and participates in transcription initiation.
(UniProt, P54269)
Summary (Interactive Fly)
Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\caup or the JBrowse view of Dmel\caup 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 5.45
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0075909
4317
693
Additional Transcript Data and Comments
Reported size (kB)
4.7, 3.4, 1.3 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0075641
73.7
693
6.99
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)
Comments
External Data
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\caup 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 (11 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
Biological Process (7 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
Cellular Component (1 term)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from sequence or structural similarity with HGNC:8632
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
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
When tin expression becomes restricted to dorsal and ventral clusters that represent cardiac and visceral mesodermal cells at stage 11, there is a stronger expression of caup mRNA around the ventral clusters of tin-positive cells.
caup transcript is expressed in both male and female genital discs of third instar larvae. Expression is more extensive and at a higher level in male than in female discs.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Mirzoyan and Pandur (FBrf0222881) use an antibody which detects both ara and caup proteins, which have similar expression patterns (see transcript expression for ara and caup). During stage 13, single ara/caup-expressing cells form alongside the row of Mef2-positive myocardial cells, and are between the eve-positive pericardial cells by stage 15.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\caup 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
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 10 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 16 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of caup
Transgenic constructs containing regulatory region of caup
Deletions and Duplications ( 10 )
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
macrochaeta & scutum
scutum & microchaeta | medial, with Scer\GAL4sca-537.4
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (16)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
9 of 15
Yes
Yes
7 of 15
No
Yes
6 of 15
No
Yes
6 of 15
No
Yes
6 of 15
No
No
5 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
 
1 of 15
No
No
 
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (16)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
7 of 15
Yes
Yes
7 of 15
Yes
No
7 of 15
Yes
Yes
6 of 15
No
Yes
6 of 15
No
Yes
6 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Rattus norvegicus (Norway rat) (14)
8 of 13
Yes
Yes
8 of 13
Yes
Yes
6 of 13
No
Yes
5 of 13
No
Yes
5 of 13
No
No
4 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (12)
6 of 12
Yes
Yes
5 of 12
No
No
5 of 12
No
Yes
5 of 12
No
No
4 of 12
No
Yes
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
Yes
1 of 12
No
No
1 of 12
No
No
Danio rerio (Zebrafish) (20)
7 of 15
Yes
Yes
7 of 15
Yes
No
6 of 15
No
Yes
6 of 15
No
Yes
6 of 15
No
No
6 of 15
No
No
5 of 15
No
Yes
5 of 15
No
No
4 of 15
No
Yes
4 of 15
No
Yes
4 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (5)
6 of 15
Yes
No
2 of 15
No
Yes
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
Yes
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (5)
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091909N1 )
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) ( EOG09150HAQ )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Musca domestica
House fly
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Glossina morsitans
Tsetse fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0EKA )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Danaus plexippus
Monarch butterfly
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
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
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Pediculus humanus
Human body louse
Pediculus humanus
Human body louse
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X07YI )
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
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
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0N9R )
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
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (6)
7 of 10
6 of 10
3 of 10
1 of 10
1 of 10
1 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 ( 0 )
    Allele
    Disease
    Interaction
    References
    Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
     
    Disease Associations of Human Orthologs (via DIOPT v7.1 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
    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
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Gene Group - Pathway Membership (FlyBase)
    External Data
    Linkouts
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    3L
    Recombination map
    3-38
    Cytogenetic map
    Sequence location
    3L:12,609,631..12,621,798 [+]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    69D1-69D1
    Limits computationally determined from genome sequence between P{PZ}l(3)0508805088 and P{PZ}l(3)0692406924
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    69D-69D
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Location
    Left of (cM)
    Right of (cM)
    Notes
    Can be 1.5% recombination between ara and mirr, with none between ara and caup.
    Stocks and Reagents
    Stocks (10)
    Genomic Clones (26)
    cDNA Clones (28)
     

    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)
    BDGP DGC clones
    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: caup CG10605
    Source for database merge of
    Additional comments
    Other Comments
    The acquisition of lateral transverse muscle fate requires the repression mediated by ara/caup of other muscle identity genes (such as slou and vg).
    S2 cells treated with dsRNA generated against this gene show reduced phagocytosis of Candida albicans compared to untreated cells.
    Iroquois complex genes (ara, caup and mirr) are required for the specification of dorsal head structures.
    The localised expression of the three genes of the iroquois complex - IRO-C (ara, caup and mirr) specifies the identity of dorsal cells in the eye. Juxtaposition of IRO-C-expressing and non-expressing cells forms a straight border that promotes growth and serves as a pattern-organising centre in the eye disc.
    The genes of the Iroquois complex (ara, caup and mirr) are necessary for notum specification. The Iro-C genes establish a signalling system that appears to organize development in the notum and dorsal wing hinge (tegula and sclerite) territories.
    When the Iroquois complex is absent, notum cells are transformed into hinge (tegula and sclerite) cells.
    A member of the IRO-C (Iroquois complex).
    The genes of the IRO-C are coordinately regulated and implicated in similar developmental processes. They are regulated in the eye by the Pc-group and trx-group genes, but not by classical modifiers of position effect variegation. Ventral silencing of the whole IRO-C in the eye occurs at the level of chromatin structure in a manner similar to that of the homeotic gene complexes, perhaps by local compaction of the region into a heterochromatin-like structure involving Pc-group products.
    ara-caup expression at patches on the wing, located one at each side of the DV compartment border, is mediated by the hh signal through its induction of high levels of ci protein in anterior cells near to the AP compartment border. High levels of ci activate dpp expression and together ci and dpp positively control ara-caup expression. The posterior border of the patches is defined by repression by en. wg accumulation at the DV border sets, also by repression, the gap between the two patches.
    The caup gene maps to the Iroquois complex and contributes to the prepattern that regulates expression of ac-sc complex genes and rho. caup is a positive, direct regulator of the ac-sc genes.
    Gene titration method has been used to search for genes involved in the determination of sense organs. A mutation of ara and caup (formerly iro) demonstrates an interaction with Df(4)M101-62f, a chromosome known to alter the development of the PNS.
    In mutants for the Iroquois gene complex, two lateral bands of the notum are completely devoid of sense organs.
    Origin and Etymology
    Discoverer
    Etymology
    The "caup" gene is named after a hero of the Araucanian American-Indian tribe.
    The 'Iroquois' gene complex is named after the native American Iroquois tribe, also called Mohawks, which shaved all but a medial stripe of hairs on the head.
    "Iroquois" is the name given to the gene complex by analogy with the American tribe that have a haircut ascribed to the Mohawk nation.
    Identification
    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/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
    Other crossreferences
    BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
    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
    Flygut - An atlas of the Drosophila adult midgut
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
    KEGG Genes - Molecular building blocks of life in the genomic space.
    modMine - A data warehouse for the modENCODE project
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DPiM - Drosophila Protein interaction map
    DroID - A comprehensive database of gene and protein interactions.
    DRSC - Results frm RNAi screens
    FLIGHT - Cell culture data for RNAi and other high-throughput technologies
    FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
    FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
    FlyMine - An integrated database for Drosophila genomics
    Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Synonyms and Secondary IDs (17)
    Reported As
    Symbol Synonym
    Secondary FlyBase IDs
    • FBgn0014195
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    References (184)