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General Information
Symbol
Dmel\cos
Species
D. melanogaster
Name
costa
Annotation Symbol
CG1708
Feature Type
FlyBase ID
FBgn0000352
Gene Model Status
Stock Availability
Enzyme Name (EC)
Adenosinetriphosphatase (3.6.1.3)
Gene Snapshot
costa (cos) encodes a kinesin-like protein and a key component in the Hedgehog (Hh) pathway that is involved in pattern formation and growth control. It forms a complex with the product of fu to restrict the activity of the Hh pathway transcription factor encoded by ci. [Date last reviewed: 2019-03-07]
Also Known As
Cos2, Costal2, cos-2, Costal-2, Costal 2
Key Links
Genomic Location
Cytogenetic map
Sequence location
2R:7,382,176..7,387,115 [-]
Recombination map
2-56
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Kinesin family. KIF27 subfamily. (O16844)
Catalytic Activity (EC)
Experimental Evidence
-
Predictions / Assertions
ATP + H(2)O = ADP + phosphate (3.6.1.3)
Summaries
Gene Group (FlyBase)
KINESINS -
Kinesin superfamily proteins (KIFs) are microtubule motor proteins which use the hydrolysis of ATP to drive directional movement along microtubules. KIFs possess a well-conserved 360 residue globular head domain which binds and hydrolyses ATP and interacts with microtubules. Many KIFs homodimerize via coiled-coil interactions in the stalk region. KIFs bind cargo through their variable tail regions and are involved in transporting organelles, protein complexes, mRNAs and the movement of spindles and chromosomes during cell division. (Adapted from FBrf0219884).
Pathway (FlyBase)
Negative Regulators of Hedgehog Signaling Pathway -
Negative regulators of hedgehog signaling down-regulation the pathway, resulting in the repression of transcription of hh-responsive genes.
Hedgehog Signaling Pathway Core Components -
The hedgehog signaling pathway is initiated by hedgehog (hh) ligand binding to the extracellular domain of patched receptor (ptc), leading to the derepression of smoothened (smo) activity. Activation of the atypical GPCR smo results in the accumulation of the transcriptional activator form of cubitus interruptus (ci) and the derepression/activation of hh target genes. In the absence of hh, smo is repressed by ptc and ci is processed to a truncated repressor form. (Adapted from FBrf0220683 and FBrf0231236).
Protein Function (UniProtKB)
Regulates cubitus interruptus (ci) processing by recruiting multiple kinases to promote its efficient phosphorylation. Scaffolds multiple kinases and ci into proximity to promote its hyperphosphorylation, which then targets it for SCFSlimb/proteasome-mediated processing to generate its repressor form. Hh signaling inhibits ci phosphorylation by interfering with the cos-ci-kinases complex formation. Negatively regulates hh-signaling pathways during various processes, including photoreceptor differentiation (PubMed:25639794, PubMed:27195754). May negatively regulate a hh-signaling pathway which functions in the intestinal immune response to bacterial uracil by activating the Duox-dependent production of reactive oxygen species (ROS) (PubMed:25639794).
(UniProt, O16844)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
cos: costa (P. Simpson)
Homozygotes, hemizygotes, and heteroallelic heterozygotes for class I alleles die as embryos with a normal cuticular phenotype. When derived from homozygous ovarian clones, however, abnormal deletion-duplication patterns in each segment are observed; posterior rows of abdominal denticles are replaced by mirror-image duplications of the anterior rows, including the segmental boundary, which is thereby duplicated; thoracic denticle belts missing; cos1 especially severely affected in this regard, including loss of entire segments. Heterozygotes derived from homozygous ovarian clones show slightly abnormal segmental patterning. Flies heterozygous for class I alleles or cos deficiencies and at the same time heterozygous for Cos alleles display pattern duplications of wings and halteres. Class II alleles are hemizygous lethal but homozygous viable. They display wing and haltere duplications indistinguishable from those found for cos1/Cos1 heterozygotes as described by Whittle. Flies heterozygous for class III alleles and Cos show moderately reduced viability and occasional wing duplications; flies homozygous for class III alleles show reduced viability and have wild-type phenotype, but enhance the pattern duplications associated with Cos/+; hemizygotes for class III alleles or heterozygotes with class I alleles are lethal or nearly so in the presence of Cos, the survivors invariably exhibiting pattern duplications of wings and halteres.
Summary (Interactive Fly)
Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\cos or the JBrowse view of Dmel\cos 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 5.48
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0089015
4763
1201
FBtr0336916
4416
1201
Additional Transcript Data and Comments
Reported size (kB)
4.9 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0088087
133.1
1201
5.64
FBpp0307854
133.1
1201
5.64
Polypeptides with Identical Sequences

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

1201 aa isoforms: cos-PA, cos-PB
Additional Polypeptide Data and Comments
Reported size (kDa)
1201 (aa); 175 (kD observed); 133 (kD predicted)
Comments
External Data
Subunit Structure (UniProtKB)
Homodimer (Potential). Binds microtubules. Interacts with ci, smo, sgg, CkIalpha and protein kinase A catalytic subunit (PubMed:15691767, PubMed:9244298). Interacts (via kinesin motor domain) with Ubr3 (PubMed:27195754).
(UniProt, O16844)
Post Translational Modification
Polyubiquitinated by Ubr3, which leads to proteasomal degradation.
(UniProt, O16844)
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\cos 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 (28 terms)
Molecular Function (11 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from physical interaction with FLYBASE:Sxl; FB:FBgn0264270
inferred from physical interaction with UniProtKB:P18431
(assigned by UniProt )
inferred from physical interaction with UniProtKB:P54367
(assigned by UniProt )
inferred from physical interaction with UniProtKB:P91682
(assigned by UniProt )
inferred from physical interaction with FLYBASE:fu; FB:FBgn0001079
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR001752
(assigned by InterPro )
inferred from biological aspect of ancestor with PANTHER:PTN000648413
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000648413
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000648413
(assigned by GO_Central )
inferred from sequence or structural similarity with FLYBASE:Khc; FB:FBgn0001308
Biological Process (10 terms)
Terms Based on Experimental Evidence (9 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000648413
(assigned by GO_Central )
Cellular Component (7 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from physical interaction with FLYBASE:fu; FB:FBgn0001079
inferred from physical interaction with FLYBASE:ci; FB:FBgn0004859
inferred from direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from sequence or structural similarity with FLYBASE:Khc; FB:FBgn0001308
inferred from biological aspect of ancestor with PANTHER:PTN000648413
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000648413
(assigned by GO_Central )
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
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
cos transcripts are detected at high levels during the first 4 hours of embryogenesis, at moderate levels between 8 and 12 hours and at low levels in later embryos. They are also detected in third instar larvae on northern blots.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
In syncytial embryos, cos protein is distributed uniformly in the cortical cytoplasm. In the late syncytial embryo, it accumulates between and apical to nuclei. From the surface, the pattern looks like a honeycomb. Throughout cellularization, cos protein is associated with furrow canals and during late cellularization is associated with expanded furrow canals. In cellular blastoderm embryos and during gastrulation, cos protein is in the cytoplasm and at the periphery of all cells. A striped pattern of cos protein is observed beginning in late stage 9 and decaying in stage 12. Each stripe is continuous along the dorsal/ventral axis both in the ectoderm and the underlying mesoderm. The stripes appear to form in the anterior compartment of each segment. In wing discs, cos protein levels are elevated in the anterior compartment
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
inferred from physical interaction with FLYBASE:fu; FB:FBgn0001079
inferred from physical interaction with FLYBASE:ci; FB:FBgn0004859
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\cos 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 ( 31 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 46 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of cos
Transgenic constructs containing regulatory region of cos
Deletions and Duplications ( 48 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
4 of 15
Yes
Yes
 
2 of 15
No
Yes
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
4 of 15
Yes
Yes
2 of 15
No
Yes
Rattus norvegicus (Norway rat) (2)
3 of 13
Yes
Yes
2 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (2)
2 of 12
Yes
Yes
1 of 12
No
Yes
Danio rerio (Zebrafish) (1)
3 of 15
Yes
Yes
Caenorhabditis elegans (Nematode, roundworm) (0)
No records found.
Arabidopsis thaliana (thale-cress) (1)
1 of 9
Yes
Yes
Saccharomyces cerevisiae (Brewer's yeast) (2)
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) ( EOG091901C2 )
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) ( EOG091500V1 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Musca domestica
House 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) ( EOG090W01OA )
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
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( None identified )
No non-Insect Arthropod orthologies identified
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
No non-Arthropod Metazoa orthologies identified
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (25)
2 of 10
2 of 10
2 of 10
2 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 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 ( 2 )
    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
    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
    RNA-RNA
    Physical Interaction
    Assay
    References
    protein-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
    suppressible
    suppressible
    External Data
    Subunit Structure (UniProtKB)
    Homodimer (Potential). Binds microtubules. Interacts with ci, smo, sgg, CkIalpha and protein kinase A catalytic subunit (PubMed:15691767, PubMed:9244298). Interacts (via kinesin motor domain) with Ubr3 (PubMed:27195754).
    (UniProt, O16844 )
    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 (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Gene Group - Pathway Membership (FlyBase)
    Negative Regulators of Hedgehog Signaling Pathway -
    Negative regulators of hedgehog signaling down-regulation the pathway, resulting in the repression of transcription of hh-responsive genes.
    Hedgehog Signaling Pathway Core Components -
    The hedgehog signaling pathway is initiated by hedgehog (hh) ligand binding to the extracellular domain of patched receptor (ptc), leading to the derepression of smoothened (smo) activity. Activation of the atypical GPCR smo results in the accumulation of the transcriptional activator form of cubitus interruptus (ci) and the derepression/activation of hh target genes. In the absence of hh, smo is repressed by ptc and ci is processed to a truncated repressor form. (Adapted from FBrf0220683 and FBrf0231236).
    External Data
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2R
    Recombination map
    2-56
    Cytogenetic map
    Sequence location
    2R:7,382,176..7,387,115 [-]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    43B1-43B2
    Limits computationally determined from genome sequence between P{lacW}vimark16722 and P{lacW}cosk16101
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    43B1-43B2
    (determined by in situ hybridisation)
    43A1-43C2
    (determined by in situ hybridisation)
    43B-43B
    (determined by in situ hybridisation)
    Placed in 42E--43E by deficiency mapping (details unspecified).
    Experimentally Determined Recombination Data
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (21)
    Genomic Clones (17)
     

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

    cDNA Clones (31)
     

    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
     
    Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for identity of: cos CG1708
    Source for database merge of
    Additional comments
    Other Comments
    cos blocks smo phosphorylation through binding with smo.
    Identified as a component of the hh signalling pathway in a genome-wide RNAi screen. dsRNA made from templates generated with primers directed affects the extent of expression of a hh signaling reporter construct in Clone 8 cells.
    dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.
    dsRNA made from templates generated with primers against this gene tested in an RNAi assay in S2 cells.
    Epistatic analysis places cos function downstream of ptc and smo.
    The last 301 amino acids of the cos COOH terminus are sufficient to provide full association with smo.
    1 allele of cos been recovered in a screen for mutations with mutant phenotypes in clones in the wing.
    fu protein phosphorylates cos protein. The primary site of phosphorylation is Ser-572, with Ser-931 being phosphorylated to a lesser extent.
    Mutants are isolated in a screen of the second chromosome isolating disc morphology defects.
    hh elicits signal transduction via a complex that includes the products of the fu, ci and cos genes. The complex binds with high affinity to microtubules in the absence of hh protein, but not when hh is present. The complex may facilitate signalling from hh by governing access of the ci product to the nucleus.
    cos encodes a kinesin-related protein that accumulates preferentially in cells capable of responding to hh signal. The product is cytoplasmic and binds microtubules. The ci protein associates with the cos product in a large protein complex, suggesting that cos directly controls the activity of ci.
    The small lobe of fu may play a role in generating the neomorphic cos phenotype displayed by an unregulated fu protein in a Su(fu)- background.
    cos function is required maternally for the patterning of the central portion of each segment, in the absence of cos activity the central portion is replaced by a mirror image duplication of the anterior partion. The cos protein may be involved in localisation or transport of other segment polarity gene products within the cell, facilitating cell polarisation.
    Viable mutations in the segmentation genes cos cause specific alterations in dpp expression within the anterior-posterior compartment boundary of the wing disc.
    Mutants display hyperplastic phenotype, imaginal disc overgrowth.
    cos negatively regulates ptc and wg transcription.
    Genetic and developmental characteristics of the imaginal disc overgrowth mutant have been determined.
    wg and en expression patterns are studied in all known segment polarity mutants to investigate the requirement of other segment polarity genes in mediating the maintenance of wg and en.
    cos suppresses the segment polarity and zygotic phenotype of one class of fu alleles and causes a lethal interaction with the other class of fu alleles.
    cos falls into the segment polarity class of genes, and has both a maternal and a zygotic action.
    Leaky alleles of cos display wing duplications that are part of a larger syndrome of polarity defects that affect the embryonic and imaginal segment. The dominant phenotype seen in flies heterozygous for the neomorphic "Cos" mutants is the result of a reduced level of function at the cos locus. Haploidy for cos enhances, whereas triploidy for cos suppresses the phenotype of "Cos" heterozygotes.
    Homozygotes, hemizygotes and heteroallelic heterozygotes for class I alleles die as embryos with a normal cuticular phenotype. When derived from homozygous ovarian clones, however, abnormal deletion-duplication patterns in each segment are observed; posterior rows of abdominal denticles are replaced by mirror-image duplications of the anterior rows, including the segmental boundary, which is thereby duplicated; thoracic denticle belts missing; cos1 especially severely affected in this regard, including loss of entire segments. Heterozygotes derived from homozygous ovarian clones show slightly abnormal segmental patterning. Flies heterozygous for class I alleles or cos deficiencies and at the same time heterozygous for Cos alleles display pattern duplications of wings and halteres. Class II alleles are hemizygous lethal but homozygous viable. They display wing and haltere duplications indistinguishable from those found for cos1/Pka-C1Cos-1 heterozygotes as described by Whittle. Flies heterozygous for class III alleles and Cos show moderately reduced viability and occasional wing duplications; flies homozygous for class III alleles show reduced viability and have wild-type phenotype, but enhance the pattern duplications associated with Cos/+; hemizygotes for class III alleles or heterozygotes with class I alleles are lethal or nearly so in the presence of Cos, the survivors invariably exhibiting pattern duplications of wings and halteres.
    Origin and Etymology
    Discoverer
    Etymology
    Identification
    External Crossreferences and Linkouts ( 64 )
    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
    UniProt/TrEMBL - Automatically annotated and unreviewed 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
    Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
    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.
    KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
    modMine - A data warehouse for the modENCODE project
    SignaLink - A signaling pathway resource with multi-layered regulatory networks.
    Linkouts
    ApoDroso - Functional genomic database for photoreceptor development, survival and function
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    DRSC - Results frm RNAi screens
    Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
    FLIGHT - Cell culture data for RNAi and other high-throughput technologies
    FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
    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.
    KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Synonyms and Secondary IDs (29)
    Reported As
    Symbol Synonym
    Cos2
    (Giordano et al., 2018, Chen et al., 2017, Zhang et al., 2017, Zhao et al., 2017, Giordani et al., 2016, Lee et al., 2016, Oh et al., 2015, Zhou et al., 2015, Andlauer et al., 2014, Kim et al., 2014, Kuzhandaivel et al., 2014, Li et al., 2014, Maier et al., 2014, Ranieri et al., 2014, Briscoe and Thérond, 2013, Chen and Jiang, 2013, Gao et al., 2013, Gao et al., 2013, Tran et al., 2013, Carroll et al., 2012, Cheng et al., 2012, Fan et al., 2012, Robbins et al., 2012, Wu et al., 2012, Marks and Kalderon, 2011, Shi et al., 2011, Zhang et al., 2011, Zhou and Kalderon, 2011, Raisin et al., 2010, Zhou and Kalderon, 2010, Farzan et al., 2009, Jia et al., 2009, Liu et al., 2008, Ogden et al., 2008, Wang and Price, 2008, Zhao and Jiang, 2008, Liu et al., 2007, Walthall et al., 2007, Zhao et al., 2007, Casso et al., 2006, De Rivoyre et al., 2006, Huangfu and Anderson, 2006, Ogden et al., 2006, Ogden et al., 2006, Osterlund and Kogerman, 2006, Singla, 2006, Varjosalo et al., 2006, Briscoe and Therond, 2005, Ho et al., 2005, Ishii, 2005, Nybakken et al., 2005, Ruel et al., 2005, Torroja et al., 2005, Walthall et al., 2005, Zhang et al., 2005, Beachy et al., 2004, Bijlsma et al., 2004, Jia et al., 2004, Kalderon, 2004, Lum and Beachy, 2004, Lum et al., 2004, Ogden et al., 2004, Stegman et al., 2004, Wang and Jiang, 2004, Zhang et al., 2004, Collins and Cohen, 2003, Fouix et al., 2003, Horabin, 2003, Jia et al., 2003, Lefers and Holmgren, 2003, Lum, 2003, Lum et al., 2003, Ogden et al., 2003, Ou et al., 2003, Ruel et al., 2003, Ruel et al., 2003, Ascano et al., 2002, Jiang, 2002, Kalderon, 2002, Lefers et al., 2002, Monnier et al., 2002, Nybakken and Perrimon, 2002, Nybakken et al., 2002, Ascano et al., 2001, Lefers et al., 2001, Miki et al., 2001, Ramirez-Weber et al., 2001, Taipale and Beachy, 2001, Vied et al., 2001, Wang et al., 2001, Stegman et al., 2000, Stegman et al., 2000, Wang et al., 2000, Chen et al., 1999, Maniatis, 1999, Marsh and Theisen, 1999, Matise and Joyner, 1999, Lee, 1998, Cadigan and Nusse, 1997, Perrimon, 1996)
    l(2)cos2
    l(2)k16101
    l(2)k16128
    Name Synonyms
    Enhancer of Epaulette
    Secondary FlyBase IDs
    • FBgn0014275
    • FBgn0021829
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    References (312)