Open Close
General Information
Symbol
Dmel\ey
Species
D. melanogaster
Name
eyeless
Annotation Symbol
CG1464
Feature Type
FlyBase ID
FBgn0005558
Gene Model Status
Stock Availability
Gene Snapshot
eyeless (ey) encodes a transcription factor that is involved in eye development as part of the retinal determination gene network, and contributes to brain formation. [Date last reviewed: 2019-03-07]
Also Known As
Pax6, l(4)33, Pax-6, eye
Key Links
Genomic Location
Cytogenetic map
Sequence location
4:697,689..721,173 [+]
Recombination map
4-0
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 paired homeobox family. (O18381)
Summaries
Gene Group (FlyBase)
PAIRED HOMEOBOX TRANSCRIPTION FACTORS -
Paired homeobox transcription factors are sequence-specific DNA binding proteins that regulate transcription. They have a characteristic paired domain in association with the homeodomain. (Adapted from FBrf0135231 and PMID:1672661).
Protein Function (UniProtKB)
Involved in eye morphogenesis.
(UniProt, O18381)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
ey: eyeless
thumb
ey4: eyeless-4
Edith M. Wallace, unpublished.
Eye size variably reduced depending on allele (see table); expressivity more variable for some alleles than for others. Tetragonal packing of facets and face-centered tetragonal bristle lattice (eyR) in place of hexagonal array of wild type [Hartman and Hayes, 1971, J. Hered. 62: 41-43 (fig.)]; associated with a failure of the horizontal secondary pigment cell to expand to give rise to the horizontal boundaries between ommatidia [Ready, Hanson, and Benzer, 1976, Dev. Biol. 53: 217-40 (fig.)]. Some ey2 flies show duplications of antennae or antennal segments with or without duplication of aristae; extra maxillary structures also observed (Shatouri, 1963, Caryologia 16: 431-37). Optical disks reduced in size [(ey1) Richards and Farrow, 1922, Proc. Oklahoma Acad. Sci. 2: 41-45; (ey2) Medvedev, 1935, Z. Indukt. Abstamm. Vererbungsl. 70: 55-72 (fig.); 1935, Tr. Inst. Genet. Akad. Nauk SSSR 10: 119-51; Steinberg, 1944, Proc. Nat. Acad. Sci. USA 30: 5-13; (ey4) Chen, 1929, J. Morphol. 47: 135-99]. Degenerating cells abundantly observed in the optic disks of third-instar larvae of ey2 [Fristrom, 1969, Mol. Gen. Genet. 103: 363-79 (fig.); Ransom, 1979, J. Embryol. Exp. Morphol. 53: 225-35]. Expressivity sensitive to genetic background [(ey4) Spofford, 1956, Genetics 41: 938-59; (ey1, ey2, ey4, eyK) Hunt and Burnet, 1969, Genet. Res. 13: 251-65]. Phenotype also responds to developmental temperature, larval density, and composition of medium. Eye size reported to increase with increased temperature in e1 (Baron, 1935, J. Exp. Zool. 70: 461-90) and eyK (Sang and Burnet, 1963, Genetics 48: 1683-99) but to decrease in eyW (Meyer, 1959, DIS 33: 97). Phenotype less extreme in flies raised under crowded conditions at 18 but not 25 (Sang and Burnet, 1963; see also Chester, 1971, DIS 46: 62-63). Eye size of four alleles increased by cholesterol deprivation and decreased by dietary deficiencies in thiamine or RNA (Hunt and Burnet, 1969). Larval feeding of lactamide to ey2 causes decreased eye size, which is of opposite sign from its effect on B (Grant and Rapport, 1980, DIS 55: 53); no such effect of lactamide on eyK observed by Sang and Burnet (1963). ey2 flies exhibit normal visual orientation in Y maze (Bulthoff, 1982, DIS 58: 31). ey2, ey4, and eyK in combination with eyg (3-35.5) results in almost complete curtailment of eye development and synthetic lethality, with the major lethal crisis at the end of the pupal stage and a minor lethal phase at pupation; rare surviving adults have brain in anterior thorax [Hunt, 1970, Genet. Res. 15: 29-34 (fig.)].
eyD: eyeless-Dominant
thumb
eyD: eyeless-Dominant
Left: head. Right: first pair of legs. From Patterson and Muller, 1930, Genetics 15: 495-577.
Eyes of heterozygotes small, outline irregular, displaced toward top and rear. Head large, often with duplicated antennae or ocelli. Basitarsus broadened distally and incompletely separated from second tarsal segment owing to interruptions of the intersegmental membrane. Polarity of bract-bristle arrangement locally reversed in regions of membrane gaps [Poodry and Schneiderman, 1976, Wilhelm Roux's Arch. Dev. Biol. 180: 175-88 (fig.)]. 27-48 sex-comb teeth disposed in more-or-less parallel longitudinal rows in males; number of transverse-bristle rows increased in females (Stern and Tokunaga, 1967, Proc. Nat. Acad. Sci. USA 57: 658-64). Extra leg joints tend to form as mirror-image duplications proximal to the normal joint between the first and second tarsal joints (Held, Duarte, and Derakhshanian, 1986, Wilhelm Roux's Arch. Dev. Biol. 195: 145-57). Clones of ey+ tissue in eyD/+ background exhibit eyD/+ phenotype (Stern and Tokunaga, 1967), but both ey+ leg disks transplanted into eyD hosts and the reciprocal transplant develop autonomously (Tokunaga, 1970, Dev. Biol. 18: 401-13). fj eyD flies have but three tarsal joints (Postlethwaite and Schneiderman, 1975, Annu. Rev. Genet. 7: 381-433). Fully dominant in triplo-4 flies (Sturtevant, 1936, Genetics 21: 448). Eye size of B; eyD/+ males larger than of B alone. Produces extreme phenotype in combination with D. D/+; eyD/+ almost completely lethal (Sobels, Kruijt, and Spronk, 1951, DIS 25: 128). Homozygous lethal; two lethal crises, one during first or second larval instar and the other just prior to or during pupal stage. Cell degeneration observed in optic disks of homozygous second-instar larval (Ransom, 1979, J. Embryol. Exp. Morphol. 53: 225-35). Larvae which are unable to pupate rescuable by injection of α-ecdysone (Arking, 1969, J. Exp. Zool. 171: 285-96). Homozygotes reaching pupal stage lack adult derivatives of eye-antennal disks; adult derivatives are formed by eyD/eyD eye-antennal disks transplanted into wildtype hosts; brain present but number of cortical cells severely reduced (Arking, Putnam, and Schubiger, 1975, J. Expt. Zool. 193: 301-12). RK2.
Summary (Interactive Fly)
transcription factor - homeodomain & pax domain - plays a crucial role in eye and brain development - controls the production of glial cells in the visual center - a temporal network transcription factor that controls neural fate of optic lobe medulla neuroblasts - , and control development of the mushroom bodies, the associative learning centers in the Drosophila brain
Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
4

Please see the GBrowse view of Dmel\ey or the JBrowse view of Dmel\ey 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 5.47
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0089236
3282
838
FBtr0089235
2854
624
FBtr0100395
2902
857
FBtr0100396
3462
898
Additional Transcript Data and Comments
Reported size (kB)
2.8 (longest cDNA)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0088300
87.6
838
7.79
FBpp0088299
65.1
624
7.01
FBpp0099809
89.4
857
7.56
FBpp0099810
93.8
898
7.38
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)
857, 838 (aa); 82.5 (kD predicted)
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\ey 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 (25 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:so; FB:FBgn0003460
inferred from physical interaction with UniProtKB:Q9VBW6
(assigned by UniProt )
inferred from physical interaction with UniProtKB:Q9VBW9
(assigned by UniProt )
inferred from mutant phenotype
Terms Based on Predictions or Assertions (0 terms)
Biological Process (19 terms)
Terms Based on Experimental Evidence (18 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
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
traceable author statement
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
expression microarray
Stage
Tissue/Position (including subcellular localization)
Reference
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
RT-PCR
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Expression in procephalic neuroblasts stage 9-11: tritocerebrum - d1, d2; deuterocerebrum - d4, v6, v7; protocerebrum - ad7, cd2, cd14, cv6, cv7, cv9, pd12, pd18, pd20, pv2
Developmental northern blots show that transcript can be detected throughout development.
ey is expressed in a cluster of approximately 10-12 cells beginning at stage 9 of embryogenesis, which correspond to the mushroom body neuroblasts. As the mushroom body neuroblasts delaminate they maintain ey expression, whereas expression diminishes in ectodermal cells that stay on the surface. ey is also expressed in a small group of neuroblasts in the deutocerebrum and tritocerebrum and in segmentally reiterated groups of three SII neuroblasts in the ventral nerve cord. Levels of expression are noted to decline by embryogenesis stage 13.
ey is expressed in mushroom body neuroblasts and in the eye disc primordium at embryonic stage 10.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
distribution deduced from reporter
Stage
Tissue/Position (including subcellular localization)
Reference
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
ey-protein is expressed in the developing larval medulla in lateral neuroblasts adjacent to l(1)sc-positive neuroepithelium and gradually weakened in neuroblasts located in more medial regions. It is co-expressed with klu-protein except for the most lateral neuroblasts. ey-protein persists in ganglion mother cells and neurons of the developing medulla.
The expression of ey, ap, and Dll were compared in outer optic lobes (OPC) starting in late third instar larvae. At this stage they were expressed as three distinct cell populations. In anterior sections, the three genes are expressed a three parallele stripes of cells that represent rows of neurons that emerge from the OPC. They correspond to progeny from the youngest to oldest neuroblasts. In middle sections, Dll-positive cells are generated in the progeny of the oldest neuroblasts, with ey-positive and ap-positive cells often placed below Dll-positive (in cells that had emerged earlier from the these neuroblasts). By the beginning of pupation, the number of cells origination from the OPC increased. A major reorganization of optic lobe structure occurs around P20 such that the three stripes are no longer distinguishable and the three cell populations are extensively interspersed within the adult medulla cortex.
ey protein is first detected in the CNS at embryonic stage 10 and corresponds to the downregulation of ind. The expression domains of ey and ind are mutually exclusive during stages 10 and 11. ey is primarily expressed in row 3-5 neuroblasts at late stage 10 and is not detected in en-positive neuroblasts.
Expression in procephalic neuroblasts stage 9-11: tritocerebrum - d1, d2; deuterocerebrum - d4, v6, v7; protocerebrum - ad7, cd2, cd14, cv6, cv7, cv9, pd12, pd18, pd20, pv2
ey protein is expressed in all cells of the second instar eye disc. By late third instar, ey protein is expressed in the eye disc anterior to the morphogenetic furrow, but not in the anterior-most part of the eye disc, where it contacts the antennal disc.
Marker for
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
Reporter: P{ey1x-GAL4.Exel}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{ey3.5-GAL4.Exel}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{ey-GAL4.S}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{ey-lacZ.C}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{ey-lacZ.mut}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GAL4-ey.H}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GawB}OK107
Stage
Tissue/Position (including subcellular localization)
Reference
eye

Comment: faint expression

ellipsoid body

Comment: faint expression

High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\ey 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 ( 45 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 40 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of ey
Transgenic constructs containing regulatory region of ey
Deletions and Duplications ( 20 )
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
antenna & eye | ectopic, with Scer\GAL4upd1-E132
antennal disc & eye | ectopic, with Scer\GAL4dpp.blk1
eye (with eyD)
eye photoreceptor cell & dorsal mesothoracic disc | ectopic, with Scer\GAL4dpp.blk1
eye photoreceptor cell & ventral thoracic disc | ectopic, with Scer\GAL4dpp.blk1
eye photoreceptor cell | ectopic & antennal disc, with Scer\GAL4dpp.blk1
haltere & eye | ectopic, with Scer\GAL4upd1-E132
haltere & ommatidium | ectopic, with Scer\GAL4dpp.blk1
leg & eye | ectopic, with Scer\GAL4dpp.blk1
leg & eye | ectopic, with Scer\GAL4upd1-E132
mushroom body & neuron
neuropil & mushroom body & pupa
proboscis & ommatidium | ectopic, with Scer\GAL4dpp.blk1
ventral thoracic disc & eye | ectopic, with Scer\GAL4dpp.blk1
wing & eye | ectopic, with Scer\GAL4MS941
wing & eye | ectopic, with Scer\GAL4upd1-E132
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (28)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
7 of 15
No
Yes
 
3 of 15
Yes
No
2 of 15
No
No
 
2 of 15
No
No
 
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
Yes
 
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
 
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
 
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) (27)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
7 of 15
No
Yes
 
4 of 15
Yes
No
2 of 15
No
No
 
2 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
Yes
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
1 of 15
Yes
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
1 of 15
No
No
Rattus norvegicus (Norway rat) (9)
4 of 13
Yes
Yes
3 of 13
No
No
2 of 13
No
No
2 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (16)
3 of 12
Yes
No
1 of 12
No
Yes
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
Yes
1 of 12
No
Yes
1 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
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
Yes
Danio rerio (Zebrafish) (30)
7 of 15
No
Yes
7 of 15
Yes
No
3 of 15
No
Yes
2 of 15
No
Yes
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
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
Yes
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
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
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (14)
9 of 15
Yes
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
Yes
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
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
Yes
Schizosaccharomyces pombe (Fission yeast) (3)
1 of 12
Yes
No
1 of 12
Yes
No
1 of 12
Yes
Yes
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091903FG )
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) ( EOG091503QS )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Mayetiola destructor
Hessian fly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( None identified )
No non-Dipteran orthologies identified
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) ( EOG091G0S4E )
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
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
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 v7.1)
Drosophila melanogaster (Fruit fly) (16)
7 of 10
4 of 10
4 of 10
4 of 10
4 of 10
3 of 10
3 of 10
3 of 10
3 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 ( 7 )
    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
    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
    suppressible
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    enhanceable
    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 (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Gene Group - Pathway Membership (FlyBase)
    External Data
    Linkouts
    SignaLink - A signaling pathway resource with multi-layered regulatory networks.
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    4
    Recombination map
    4-0
    Cytogenetic map
    Sequence location
    4:697,689..721,173 [+]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    102D4-102D5
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    Experimentally Determined Recombination Data
    Location
    Left of (cM)
    Right of (cM)
    Notes
    Mapped by recombination in diplo-4 triploids.
    Stocks and Reagents
    Stocks (88)
    Genomic Clones (23)
    cDNA Clones (14)
     

    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
         
        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: ey CG1464
        Source for database merge of
        Additional comments
        ey2 is the most frequently used allele. Four lethal alleles, formerly l(4)10 = l(4)33, assigned to the ey locus by Hochman (1971); they produce a low incidence of escapers with reduced eyes and but partially complement eyR.
        Other Comments
        ey induces expression of dac.
        The DNA binding activity of ey relies on the PAI and L regions of the PD domain.
        pb and ey have antagonistic roles in head development.
        ey collaborates with hh and dpp signalling in eye induction.
        gish, so, ey and hh act in the posterior region of the eye disc to prevent precocious glial cell migration.
        ey is not responsible for the development of head structures derived from the antennal disc, but is primarily required to inhibit cell death and to promote eye development.
        Overexpression of ey produces a phenotype in the embryonic nervous system.
        ey has a function in the structural development of the mushroom body.
        ey and dac may have a role in axon pathway selection during embryogenesis.
        ey and dac do not seem to regulate each other's expression in the developing mushroom body.
        A single amino acid substitution in the ey protein is able to change the DNA-binding affinity of its paired domain to that of toy.
        toy acts upstream of ey in the control of eye development. Despite the similarity in structure and expression of toy and ey, normal expression of toy cannot compensate for the loss of ey function.
        ey protein activates transcription of so by direct interaction with an eye-specific enhancer in the long intron of so.
        so and eya are two mediators of the eye inducing activity of ey. ey appears to induce the initial expression of so and eya in the eye disc. so and eya then participate in a positive feedback loop that regulates the expression of all three genes. In the embryonic head, however, so acts in parallel to ey and toy. The epistatic relationships among the corresponding vertebrate homologs are very similar to those observed in Drosophila.
        tsh and ey induce the expression of each other, acting in a gene network that functions to specify eye identity.
        eya is essential for ey function.
        dac and ey induce the expression of each other and dac is required for ectopic retinal development driven by ey misexpression. Results suggest dac and ey are likely to function together in the control of retinal cell-fate specification at the early stages of eye development.
        The pattern of ey expression during eye and Bolwig organ development has been analysed. ey directly regulates ninaE expression in photoreceptor cells.
        Used in an investigation to address the relationship between retrotransposons and retroviruses and the coadaptation of these retroelements to their host genomes. Results indicate retrotransposons are heterogeneous in contrast to retroviruses, suggesting different modes of evolution by slippage-like mechanisms.
        Targeted expression of ey in various imaginal disc primordia results in ectopic eyes on the wings, legs and antennae. The ectopic eyes appear morphologically normal and consist of groups of fully differentiated ommatidia with a complete set of photoreceptor cells. Similar ectopic eyes can be induced by targeted expression of the Mmus\Pax6 gene.
        Comparisons of early development to that in other insects have revealed conservation of some aspects of development, as well as differences that may explain variations in early patterning events.
        Eye size variably reduced depending on allele (see table); expressivity more variable for some alleles than for others. Tetragonal packing of facets and face-centered tetragonal bristle lattice (eyR) in place of hexagonal array of wild type (Hartman and Hayes, 1971); associated with a failure of the horizontal secondary pigment cell to expand to give rise to the horizontal boundaries between ommatidia (Ready, Hanson and Benzer, 1976). Some ey2 flies show duplications of antennae or antennal segments with or without duplication of aristae; extra maxillary structures also observed (Shatouri, 1963). Optical discs reduced in size (ey<up>1) Richards and Farrow, 1922; (ey2) Medvedev, 1935; Medvedev, 1935; Steinberg, 1944; (ey4) Chen, 1929</up>. Degenerating cells abundantly observed in the optic discs of third instar larvae of ey2 (Fristrom, 1969; Ransom, 1979). Expressivity sensitive to genetic background (ey<up>4) Spofford, 1956; (ey1, ey2, ey4, eyK) Hunt and Burnet, 1969</up>. Phenotype also responds to developmental temperature, larval density, and composition of medium. Eye size reported to increase with increased temperature in e1 (Baron, 1935) and eyK (Sang and Burnet, 1963) but to decrease in eyW (Meyer, 1959). Phenotype less extreme in flies raised under crowded conditions at 18oC but not 25oC (Sang and Burnet, 1963; see also Chester, 1971). Eye size of four alleles increased by cholesterol deprivation and decreased by dietary deficiencies in thiamine or RNA (Hunt and Burnet, 1969). Larval feeding of lactamide to ey2 causes decreased eye size, which is of opposite sign from its effect on B (Grant and Rapport, 1980); no such effect of lactamide on eyK observed by Sang and Burnet (1963). ey2 flies exhibit normal visual orientation in Y maze (Bulthoff, 1982). ey2, ey4 and eyK in combination with eyg (3-35.5) results in almost complete curtailment of eye development and synthetic lethality, with the major lethal crisis at the end of the pupal stage and a minor lethal phase at pupation; rare surviving adults have brain in anterior thorax (Hunt, 1970).
        Origin and Etymology
        Discoverer
        Hoge, May 1914.
        Etymology
        Identification
        External Crossreferences and Linkouts ( 79 )
        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
        GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
        KEGG Genes - Molecular building blocks of life in the genomic space.
        modMine - A data warehouse for the modENCODE project
        SignaLink - A signaling pathway resource with multi-layered regulatory networks.
        Linkouts
        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.
        MIST (genetic) - An integrated Molecular Interaction Database
        MIST (protein-protein) - An integrated Molecular Interaction Database
        Synonyms and Secondary IDs (19)
        Reported As
        Symbol Synonym
        ey
        (Curt et al., 2019, Gao et al., 2019, Meltzer et al., 2019, Papadopoulos et al., 2019, Shokri et al., 2019, Bischof et al., 2018, Davie et al., 2018, Kumar, 2018, Magri et al., 2018, Palliyil et al., 2018, Sarkar et al., 2018, Zhu et al., 2018, Aggarwal et al., 2017, Erceg et al., 2017, Hu et al., 2017.6.13, Transgenic RNAi Project members, 2017-, Zhu et al., 2017, Aggarwal et al., 2016, Beira and Paro, 2016, Bielmeier et al., 2016, Blick et al., 2016, Bürglin and Affolter, 2016, Du et al., 2016, Lin and Potter, 2016, Peng et al., 2016, Sarov et al., 2016, Weasner et al., 2016, Doggett et al., 2015, Schertel et al., 2015, Tanaka-Matakatsu et al., 2015, Ugrankar et al., 2015, Wittkorn et al., 2015, Zhao et al., 2015, Ashwal-Fluss et al., 2014, Bauerly et al., 2014, Bertet et al., 2014, Blaquiere et al., 2014, Karandikar et al., 2014, Tanaka-Matakatsu et al., 2014, Zhou et al., 2014, Atkins et al., 2013, Li et al., 2013, Marinho et al., 2013, Moran et al., 2013, Naval-Sánchez et al., 2013, Suzuki et al., 2013, Weasner and Kumar, 2013, Kunz et al., 2012, Manning et al., 2012, Najafov et al., 2012, Nfonsam et al., 2012, Spokony and White, 2012.11.14, Venables et al., 2012, Wang and Sun, 2012, Brockmann et al., 2011, Datta et al., 2011, Gehring, 2011, Kearse et al., 2011, Kopyl et al., 2011, Michaut et al., 2011, Miles et al., 2011, Morante et al., 2011, Stephan et al., 2011, Sultana et al., 2011, Aerts et al., 2010, Baig et al., 2010, Blanco et al., 2010, Kong et al., 2010, Lopes and Casares, 2010, Papadopoulos et al., 2010, Popov et al., 2010, Roignant et al., 2010, Salzer and Kumar, 2010, Salzer et al., 2010, Clements et al., 2009, Datta et al., 2009, Firth and Baker, 2009, Furukubo-Tokunaga et al., 2009, Gutierrez-Aviño et al., 2009, Jacobsson et al., 2009, Kurusu et al., 2009, Lee et al., 2009, Schulze et al., 2009, Weasner et al., 2009, Anderson and Kumar, 2008, Braid and Verheyen, 2008, Brás-Pereira and Casares, 2008, Chen et al., 2008, Clements et al., 2008, Clements et al., 2008, Duong et al., 2008, Jacobsson et al., 2008, McDermott and Kliman, 2008, Morante and Desplan, 2008, Plaza et al., 2008, Weasner and Kumar, 2008, Yao et al., 2008, Curtiss et al., 2007, Fic et al., 2007, Kumar and Anderson, 2007, Kumar and Datta, 2007, Kumar and Luhur, 2007, Kumar and Salzer, 2007, Kumar and Weanser, 2007, Luque and Milan, 2007, Sprecher et al., 2007, Von Ohlen et al., 2007, Wang et al., 2007, Weasner et al., 2007, Yoshioka et al., 2007, Zeitlinger et al., 2007, Anderson et al., 2006, Friedrich, 2006, Kobayashi et al., 2006, Oakley et al., 2006, Ostrin et al., 2006, Urbach et al., 2006, Apitz et al., 2005, Choi et al., 2005, Roederer et al., 2005, Zhang et al., 2005, Podemski et al., 2004, Punzo et al., 2004, Noor and Kliman, 2003, Kronhamn et al., 2002, Onuma et al., 2002, Halder et al., 1998)
        l(4)102CDh
        l(4)102CDr
        Name Synonyms
        eyeless
        (Papadopoulos et al., 2019, Croset et al., 2018, Kumar, 2018, Bielmeier et al., 2016, Jin and Mardon, 2016, Nässel and Vanden Broeck, 2016, Suzuki et al., 2016, Hales et al., 2015, Nässel et al., 2015, Tanaka-Matakatsu et al., 2015, Bauerly et al., 2014, Marinho et al., 2013, Naval-Sánchez et al., 2013, Southall et al., 2013, Weasner and Kumar, 2013, Wredenberg et al., 2013, Maclellan et al., 2012, Najafov et al., 2012, Venables et al., 2012, Yang et al., 2012, Gehring, 2011, Kearse et al., 2011, Miles et al., 2011, Stephan et al., 2011, Sultana et al., 2011, Baig et al., 2010, Blanco et al., 2010, Lopes and Casares, 2010, Papadopoulos et al., 2010, Roignant et al., 2010, Salzer and Kumar, 2010, Voolstra et al., 2010, Clements et al., 2009, Furukubo-Tokunaga et al., 2009, Gutierrez-Aviño et al., 2009, Jacobsson et al., 2009, Schulze et al., 2009, Weasner et al., 2009, Chen et al., 2008, Clements et al., 2008, Clements et al., 2008, Duong et al., 2008, Jacobsson et al., 2008, Legent et al., 2008, Morante and Desplan, 2008, Wang and Sun, 2008, Fic et al., 2007, Kumar and Anderson, 2007, Kumar and Datta, 2007, Kumar and Luhur, 2007, Kumar and Salzer, 2007, Kumar and Weanser, 2007, Legent et al., 2007, Li and Baker, 2007, Luque and Milan, 2007, Pereanu et al., 2007, Schwartz and Pirrotta, 2007, Von Ohlen et al., 2007, Wang et al., 2007, Yoshioka et al., 2007, Zeitlinger et al., 2007, Fernald, 2006, Friedrich, 2006, Ostrin et al., 2006, Roederer et al., 2005, Noor and Kliman, 2003, Xu, 1998.8.25)
        tumor-head-63
        Secondary FlyBase IDs
        • FBgn0000616
        • FBgn0004525
        • FBgn0004547
        • FBgn0010156
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        References (643)