FB2025_04 , released October 2, 2025
Gene: Dmel\ex
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General Information
Symbol
Dmel\ex
Species
D. melanogaster
Name
expanded
Annotation Symbol
CG4114
Feature Type
protein_coding_gene
FlyBase ID
FBgn0004583
Gene Model Status
Current
Stock Availability
35 publicly available
Gene Summary
expanded (ex) encodes a FERM-domain containing protein that localizes to apical cell-cell junctions, where it promotes Hippo signaling by physically interacting with multiple Hippo pathway components. Through its regulation of Hippo signaling, the product of ex functions as an inhibitor of growth. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Key Links
Genomic Location
Cytogenetic map
21C2-21C2
Sequence location
Recombination map
2-0.7
RefSeq locus
NT_033779 REGION:431227..448701
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (26 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
enables DNA-binding transcription factor binding
inferred from physical interaction with UniProtKB:Q45VV3
(Hu et al., 2016)
enables kinase binding
inferred from physical interaction with UniProtKB:Q9VZI2
(Hu et al., 2016)
enables protein binding
inferred from physical interaction with FLYBASE:Mer; FB:FBgn0086384
(McCartney et al., 2000)
inferred from physical interaction with UniProtKB:Q9VFG8
(Yu et al., 2010)
enables protein sequestering activity
inferred from physical interaction with FLYBASE:yki; FB:FBgn0034970,FLYBASE:Ptp61F; FB:FBgn0267487
(Liu et al., 2022)
inferred from physical interaction with FLYBASE:yki; FB:FBgn0034970
(Badouel et al., 2009)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (17 terms)
Terms Based on Experimental Evidence (17 terms)
CV Term
Evidence
References
involved_in border follicle cell migration
inferred from mutant phenotype
(Lucas et al., 2013)
involved_in compound eye morphogenesis
inferred from mutant phenotype
(Pellock et al., 2007)
inferred from genetic interaction with FLYBASE:crb; FB:FBgn0259685
(Ling et al., 2010)
involved_in compound eye photoreceptor cell differentiation
inferred from mutant phenotype
(Pellock et al., 2007)
inferred from genetic interaction with FLYBASE:Mer; FB:FBgn0086384
(Maitra et al., 2006)
involved_in endocytic recycling
inferred from genetic interaction with FLYBASE:Mer; FB:FBgn0086384
(Maitra et al., 2006)
involved_in energy homeostasis
inferred from genetic interaction with FLYBASE:mir-278; FB:FBgn0262403
(Teleman and Cohen, 2006)
involved_in follicle cell of egg chamber development
inferred from mutant phenotype
(Jia et al., 2015)
involved_in negative regulation of cell population proliferation
inferred from mutant phenotype
(Badouel et al., 2009, Pellock et al., 2007, McCartney et al., 2000)
inferred from genetic interaction with FLYBASE:Mer; FB:FBgn0086384
(Hamaratoglu et al., 2006, McCartney et al., 2000)
involved_in negative regulation of imaginal disc growth
inferred from genetic interaction with FLYBASE:wts; FB:FBgn0011739
(Hamaratoglu et al., 2006)
inferred from mutant phenotype
(Rauskolb et al., 2011, Pellock et al., 2007)
involved_in negative regulation of organ growth
inferred from genetic interaction with FLYBASE:wts; FB:FBgn0011739
(Feng and Irvine, 2007)
inferred from genetic interaction with FLYBASE:Mer; FB:FBgn0086384
(Hamaratoglu et al., 2006)
inferred from mutant phenotype
(Pellock et al., 2007)
involved_in negative regulation of TORC1 signaling
inferred from mutant phenotype
(Liu et al., 2022)
involved_in positive regulation of apoptotic process
inferred from genetic interaction with FLYBASE:Mer; FB:FBgn0086384
(Hamaratoglu et al., 2006)
inferred from mutant phenotype
(Willecke et al., 2006)
involved_in positive regulation of hippo signaling
inferred from mutant phenotype
(Fletcher et al., 2018, Yu and Pan, 2018, Hu et al., 2016, Yang et al., 2015, Zhang et al., 2015, Pellock et al., 2007)
inferred from direct assay
(Liu et al., 2022, Yu et al., 2010, Oh et al., 2009)
inferred from genetic interaction with FLYBASE:ft; FB:FBgn0001075
(Oh and Irvine, 2008, Willecke et al., 2006)
inferred from genetic interaction with FLYBASE:hpo; FB:FBgn0261456
(Hamaratoglu et al., 2006)
inferred from genetic interaction with FLYBASE:yki; FB:FBgn0034970,FLYBASE:Mer; FB:FBgn0086384
(Hamaratoglu et al., 2009)
inferred from genetic interaction with FLYBASE:wts; FB:FBgn0011739
(Badouel et al., 2009, Feng and Irvine, 2007)
inferred from genetic interaction with FLYBASE:kibra; FB:FBgn0262127
(Baumgartner et al., 2010)
inferred from physical interaction with FLYBASE:yki; FB:FBgn0034970
(Badouel et al., 2009)
involved_in positive regulation of proteasomal ubiquitin-dependent protein catabolic process
inferred from mutant phenotype
(Ribeiro et al., 2014)
involved_in regulation of cell differentiation
inferred from genetic interaction with FLYBASE:Mer; FB:FBgn0086384
(McCartney et al., 2000)
involved_in regulation of exit from mitosis
inferred from mutant phenotype
(Pellock et al., 2007)
involved_in regulation of growth
inferred from genetic interaction with FLYBASE:RpL36; FB:FBgn0002579
(Tyler et al., 2007)
involved_in regulation of signal transduction
inferred from genetic interaction with FLYBASE:Mer; FB:FBgn0086384
(Maitra et al., 2006)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
involved_in positive regulation of hippo signaling
inferred from biological aspect of ancestor with PANTHER:PTN000334995
(GO Reference Genome Project, 2011-)
Cellular Component (5 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
located_in apicolateral plasma membrane
inferred from direct assay
(Oh et al., 2009, Feng and Irvine, 2007)
colocalizes_with cell-cell junction
inferred from direct assay
(Su et al., 2017)
located_in cytoplasmic side of apical plasma membrane
inferred from direct assay
(Ribeiro et al., 2014, Genevet et al., 2010, Ling et al., 2010, McCartney et al., 2000)
part_of Kibra-Ex-Mer complex
inferred from direct assay
(Yu et al., 2010)
inferred from physical interaction with FLYBASE:Mer; FB:FBgn0086384,FLYBASE:kibra; FB:FBgn0262127
(Genevet et al., 2010)
located_in subapical part of cell
inferred from direct assay
(Wang et al., 2019)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in cytoplasmic side of apical plasma membrane
inferred from biological aspect of ancestor with PANTHER:PTN000334995
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
expanded (ex) encodes a FERM-domain containing protein that localizes to apical cell-cell junctions, where it promotes Hippo signaling by physically interacting with multiple Hippo pathway components. Through its regulation of Hippo signaling, the product of ex functions as an inhibitor of growth. [Date last reviewed: 2019-03-07]
Pathway (FlyBase)
POSITIVE REGULATORS OF HIPPO SIGNALING PATHWAY -
The Hippo signaling pathway is an intracellular kinase cascade in which hpo kinase in complex with sav, phosphorylates wts kinase which, in turn, phosphorylates yki transcriptional co-activator leading to its cytosolic retention. Positive regulators of the pathway, enhance the cytosolic retention of yki, negatively regulating tissue growth (Adapted from FBrf0224870).
Gene Group (FlyBase)
KIBRA-EX-MER COMPLEX -
The KEM complex is an apical protein complex that contains the proteins Kibra, Expanded and Merlin. It positively regulates the Hippo pathway, acting as a scaffolding complex. (Adapted from FBrf0210017).
Protein Function (UniProtKB)
Activates the Hippo/SWH (Sav/Wts/Hpo) signaling pathway, a signaling pathway that plays a pivotal role in organ size control and tumor suppression by restricting proliferation and promoting apoptosis (PubMed:20159598, PubMed:26954546, PubMed:27462444, PubMed:8269855). The core of this pathway is composed of a kinase cascade wherein Hippo (Hpo), in complex with its regulatory protein Salvador (Sav), phosphorylates and activates Warts (Wts) in complex with its regulatory protein Mats, which in turn phosphorylates and inactivates the Yorkie (Yki) oncoprotein. Ex acts synergistically along with Mer and Kibra to regulate the Hippo signaling pathway (PubMed:20159598). Involved in the control of cell proliferation in imaginal disks (PubMed:27462444, PubMed:8269855). May bind to certain proteins of signal transduction pathways by interaction with their SH3 domains (PubMed:20159598). Required for apical localization of Schip1 (PubMed:26954546).
(UniProt, Q07436)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
brl: braille (M.P. Scott)
Recessive mutation with complete penetrance and variable expressivity. Strongest phenotypes include partial to full transformation of one or both eyes into antennae. Less severe adult phenotypes include excess bristles under the eyes, duplication of the more anterior set of scutellar bristles (macrochaetae), and a spoon-like curvature of the wings. Homozygotes are viable and fertile.
ex: expanded
thumb
ex: expanded
From Stern and Bridges, 1926, Genetics 11: 503-30.
Wings extremely wide and large, sometimes curved and divergent. Effect produced in prepupal wing, probably by influence on cell division (Waddington, 1940, J. Genet. 41: 75-139). Eyes slightly reduced in size and roughish. Body large. RK2.
Summary (Interactive Fly)

Band 4.1 protein with three potential SH3-binding sites - localization to adherens junctions allows Expanded to exert effects on multiple processes involved in disc development, including effects on cell proliferation, cell fate determination, and tissue polarity

Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\ex for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q07436)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Gene model reviewed during 5.43

Gene model reviewed during 5.52

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
ex-RA
FBtr0078059
NM_057492
6672
1427
ex-RB
FBtr0329832
NM_001272894
6454
1427
Additional Transcript Data and Comments
Reported size (kB)

6.9 (northern blot)

(Boedigheimer and Laughon, 1993)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
ex-PA
FBpp0077719
153.6
1427
8.45
Q07436
NP_476840
AAF51495
ex-PB
FBpp0302878
153.6
1427
8.45
Q07436
NP_001259823
AGB92360
Polypeptides with Identical Sequences

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

1427 aa isoforms: ex-PA, ex-PB
Additional Polypeptide Data and Comments
Reported size (kDa)

1429 (aa)

(Boedigheimer and Laughon, 1993)
Comments
External Data
Subunit Structure (UniProtKB)

Forms a complex with Kibra and Mer (PubMed:20159598). Interacts (via RXPPXY motif) with Kibra (via domain WW 1) (PubMed:20159598). Interacts with Mer and Hpo (via SARAH domain) (PubMed:20159598). Interacts with Schip1; the interaction results in recruitment of Schip1 to the apical cell membrane (PubMed:26954546). Interacts with ack and yki (PubMed:27462444).

(UniProt, Q07436)
Post Translational Modification

Phosphorylated by Ack at several tyrosines including Tyr-227, Tyr-423, Tyr-679, Tyr-766 and Tyr-1103.

(UniProt, Q07436)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

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

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

-0.72

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
larval stage
imaginal disc
(Boedigheimer and Laughon, 1993)
Additional Descriptive Data

ex transcripts are apically localized in imaginal discs.

(Boedigheimer and Laughon, 1993)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage -- larval stage
neuroblast NB5-3
(Lacin and Truman, 2016)
larval stage
leg disc
(Boedigheimer and Laughon, 1993)
wing disc
(Boedigheimer and Laughon, 1993)
imaginal disc
(Boedigheimer and Laughon, 1993)
third instar larval stage
eye disc
(Milton et al., 2010)
pupal stage P6
eye disc | faint
(Milton et al., 2010)
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
eye disc
(Milton et al., 2010)
pupal stage P6
eye disc | faint
(Milton et al., 2010)
Additional Descriptive Data

ex protein is localized to the extreme apical cell surface.

(Milton et al., 2010, Boedigheimer and Laughon, 1993)

ex protein is detected by early third instar in imaginal discs. It is fairly uniform throughout leg discs but is concentrated in the wing pouch relative to other areas of the wing disc.

(Boedigheimer and Laughon, 1993)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in apicolateral plasma membrane
inferred from direct assay
(Oh et al., 2009, Feng and Irvine, 2007)
colocalizes_with cell-cell junction
inferred from direct assay
(Su et al., 2017)
located_in cytoplasmic side of apical plasma membrane
inferred from direct assay
(Ribeiro et al., 2014, Genevet et al., 2010, Ling et al., 2010, McCartney et al., 2000)
part_of Kibra-Ex-Mer complex
inferred from direct assay
(Yu et al., 2010)
inferred from physical interaction with FLYBASE:Mer; FB:FBgn0086384,FLYBASE:kibra; FB:FBgn0262127
(Genevet et al., 2010)
located_in subapical part of cell
inferred from direct assay
(Wang et al., 2019)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\ex in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 37 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 42 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of ex
Transgenic constructs containing regulatory region of ex
Aberrations (Deficiencies and Duplications) ( 12 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal
lethal (with Df(2L)BSC107)
lethal | recessive
lethal - all die before end of P-stage | recessive
partially lethal - majority die
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4tin.CΔ4
Sterility
Allele
fertile
fertile, with Scer\GAL4VP16.nanos.UTR
Other Phenotypes
Allele
abnormal cell migration
abnormal cell migration | oogenesis | somatic clone
abnormal mitotic cell cycle
abnormal neuroanatomy | embryonic stage
abnormal planar polarity, with Scer\GAL4hs.2sev
abnormal planar polarity | somatic clone
abnormal size, with Scer\GAL4nub-AC-62
abnormal size | adult stage
abnormal size | adult stage, with Scer\GAL4en.PU
abnormal size | adult stage, with Scer\GAL4en-e16E
abnormal size | adult stage, with Scer\GAL4nub.PU
abnormal size | adult stage, with Scer\GAL4nub-AC-62
abnormal size | adult stage | somatic clone
abnormal size | larval stage, with Scer\GAL4dpp.PU
abnormal size | somatic clone | third instar larval stage
abnormal size | third instar larval stage, with Scer\GAL4hh-Gal4
decreased cell number | larval stage | somatic clone, with Scer\GAL4FRT.Act5C
decreased cell number | third instar larval stage
decreased cell size | recessive
decreased size | adult stage
decreased size | adult stage, with Scer\GAL4bbg-C96
hyperplasia
hyperplasia | larval stage (with exMGH1)
hyperplasia | larval stage (with exMGH2)
hyperplasia | recessive
hyperplasia | somatic clone
hypoplasia | adult stage, with Scer\GAL4nub.PU
increased cell death, with Scer\GAL4bi-omb-Gal4
increased cell death, with Scer\GAL4ey.PH
increased cell death, with Scer\GAL4GMR.PF
increased cell death, with Scer\GAL4hs.2sev
increased cell death | larval stage, with Scer\GAL4en.PU
increased cell death | third instar larval stage, with Scer\GAL4GMR.PF
increased cell growth rate | somatic clone, with Scer\GAL4Act5C.CoinFLP-FRT
increased cell number
increased cell number | cell non-autonomous, with Scer\GAL4C587
increased cell number | cell non-autonomous | somatic clone
increased cell number | larval stage
increased cell number | pupal stage, with Scer\GAL4GMR.PFa
increased occurrence of cell division
increased occurrence of cell division | larval stage | somatic clone
increased size | adult stage
increased size | adult stage, with Scer\GAL4en-e16E
increased size | adult stage, with Scer\GAL4ptc.PU
increased size | adult stage | somatic clone - tissue specific
neoplasia | adult stage | cell non-autonomous | male | somatic clone
some die during pharate adult stage | recessive
uncoordinated | recessive
visible
visible, with Scer\GAL432B
visible, with Scer\GAL4bi-omb-Gal4
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4dpp.blk1
visible, with Scer\GAL4ey.PH
visible, with Scer\GAL4ey.PU
visible, with Scer\GAL4GMR.PF
visible, with Scer\GAL4GMR.PU
visible, with Scer\GAL4hh-Gal4
visible, with Scer\GAL4hs.2sev
visible, with Scer\GAL4pnr-MD237
visible | adult stage
visible | adult stage, with Scer\GAL4bbg-C96
visible | adult stage, with Scer\GAL4Bx-MS1096
visible | adult stage, with Scer\GAL4en.PU
visible | adult stage, with Scer\GAL4en-e16E
visible | adult stage, with Scer\GAL4GMR.PF
visible | adult stage, with Scer\GAL4GMR.PU
visible | adult stage, with Scer\GAL4nub.PU
visible | adult stage, with Scer\GAL4ptc.PU
visible | adult stage | somatic clone
visible | recessive
visible | recessive | cold sensitive
visible | somatic clone
visible | somatic clone, with Scer\GAL4Act5C.CoinFLP-FRT
Phenotype manifest in
Allele
adult abdomen
adult head | cold sensitive
adult head | somatic clone
adult thorax
antenna
antenna | adult stage | somatic clone
antennal sense organ
anterior crossvein
anterior crossvein | ectopic, with Scer\GAL432B
anterior scutellar bristle
arista
border follicle cell
border follicle cell | somatic clone
cone cell, with Scer\GAL4hs.2sev
embryonic/larval optic lobe
eye
eye, with Scer\GAL432B
eye, with Scer\GAL4ey.PH
eye, with Scer\GAL4ey.PU
eye, with Scer\GAL4GMR.PF
eye, with Scer\GAL4GMR.PU
eye, with Scer\GAL4hs.2sev
eye | adult stage, with Scer\GAL4GMR.PU
eye | adult stage | somatic clone
eye | anterior, with Scer\GAL4hs.2sev
eye | somatic clone
eye | somatic clone, with Scer\GAL4Act5C.CoinFLP-FRT
eye | somatic clone - tissue specific
eye-antennal disc
eye-antennal disc (with exMGH1)
eye-antennal disc (with exMGH2)
eye-antennal disc | third instar larval stage
eye disc
eye disc, with Scer\GAL4ey.PH
eye disc | pupal stage | somatic clone
eye disc | somatic clone
eye disc | third instar larval stage
eye disc | third instar larval stage, with Scer\GAL4GMR.PF
eye disc morphogenetic furrow | third instar larval stage, with Scer\GAL4GMR.PF
eye photoreceptor cell, with Scer\GAL4hs.2sev
first posterior wing cell, with Scer\GAL4dpp.blk1
haltere disc
head
imaginal disc
interommatidial bristle & eye disc & pupa
interommatidial bristle | pupal stage | somatic clone | increased number
interommatidial bristle | somatic clone
interommatidial bristle | somatic clone | increased number
interommatidial cell | pupal stage, with Scer\GAL4GMR.PF
interommatidial cell | pupal stage | increased number, with Scer\GAL4GMR.PFa
interommatidial cell | pupal stage | somatic clone | increased number
larval glial cell | third instar larval stage
larval RP2 motor neuron | embryonic stage
leg
leg, with Scer\GAL4bi-omb-Gal4
leg, with Scer\GAL4Bx-MS1096
leg | cold sensitive
mesonotum, with Scer\GAL4pnr-MD237
mesothoracic leg disc
mesothoracic tarsal segment 3
mesothoracic tarsal segment 4
mesothoracic tarsal segment 5
metathoracic leg disc
metathoracic tarsal segment 3
metathoracic tarsal segment 4
metathoracic tarsal segment 5
ommatidium
ommatidium, with Scer\GAL432B
ommatidium, with Scer\GAL4ey.PH
ommatidium, with Scer\GAL4hs.2sev
ommatidium | cell autonomous | somatic clone
ommatidium | cell non-autonomous | somatic clone
ommatidium | pupal stage | somatic clone
ommatidium | somatic clone
photoreceptor neuron (with exMGH1)
photoreceptor neuron (with exMGH2)
photoreceptor neuron | somatic clone
pigment cell, with Scer\GAL432B
pigment cell, with Scer\GAL4hs.2sev
pigment cell | increased number
posterior crossvein
posterior crossvein, with Scer\GAL4en-e16E
postgenal trichoid sensillum
prothoracic leg disc
prothoracic tarsal segment 3
prothoracic tarsal segment 4
prothoracic tarsal segment 5
retina | pupal stage | somatic clone
spermatogonial cyst | cell non-autonomous, with Scer\GAL4C587
spermatogonial cyst | cell non-autonomous | somatic clone
testis | adult stage | cell non-autonomous | somatic clone
thorax
vibrissal bristle
wing
wing, with Scer\GAL432B
wing, with Scer\GAL4bbg-C96
wing, with Scer\GAL4bi-omb-Gal4
wing, with Scer\GAL4Bx-MS1096
wing, with Scer\GAL4en-e16E
wing, with Scer\GAL4nub.PU
wing, with Scer\GAL4nub-AC-62
wing, with Scer\GAL4ptc.PU
wing | posterior, with Scer\GAL4en-e16E
wing | prepupal stage
wing | somatic clone
wing blade, with Scer\GAL4Bx-MS1096
wing blade, with Scer\GAL4nub.PU
wing blade, with Scer\GAL4nub-AC-62
wing disc
wing disc, with Scer\GAL4dpp.PU
wing disc, with Scer\GAL4hh-Gal4
wing disc (with exMGH1)
wing disc (with exMGH2)
wing disc | larval stage
wing disc | larval stage | somatic clone, with Scer\GAL4FRT.Act5C
wing disc | somatic clone | third instar larval stage
wing disc | third instar larval stage, with Scer\GAL4hh-Gal4
wing disc | third instar larval stage (with exAP50)
wing disc | third instar larval stage (with exe1)
wing disc posterior compartment, with Scer\GAL4en.PU
wing margin, with Scer\GAL432B
wing margin, with Scer\GAL4Bx-MS1096
wing posterior compartment, with Scer\GAL4en.PU
wing vein, with Scer\GAL432B
wing vein, with Scer\GAL4bi-omb-Gal4
wing vein, with Scer\GAL4dpp.blk1
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (29)
Hsap\FRMD1
7 of 14
Yes
Yes
Hsap\FRMD6
4 of 14
No
Yes
Hsap\EPB41
1 of 14
No
No
Hsap\EPB41L1
1 of 14
No
No
Hsap\EPB41L2
1 of 14
No
No
1  
Hsap\EPB41L3
1 of 14
No
No
Hsap\EPB41L4A
1 of 14
No
No
Hsap\EPB41L4B
1 of 14
No
No
Hsap\EPB41L5
1 of 14
No
No
Hsap\EZR
1 of 14
No
No
1  
Hsap\FARP1
1 of 14
No
No
Hsap\FARP2
1 of 14
No
No
Hsap\FRMD3
1 of 14
No
No
Hsap\FRMD4A
1 of 14
No
No
Hsap\FRMD4B
1 of 14
No
No
Hsap\FRMD5
1 of 14
No
No
7  
Hsap\FRMD7
1 of 14
No
No
Hsap\FRMPD2
1 of 14
No
No
Hsap\LAMB2
1 of 14
No
No
Hsap\MSN
1 of 14
No
No
Hsap\MYLIP
1 of 14
No
No
Hsap\NF2
1 of 14
No
No
10  
Hsap\PTPN3
1 of 14
No
No
Hsap\PTPN4
1 of 14
No
No
Hsap\PTPN13
1 of 14
No
No
Hsap\PTPN14
1 of 14
No
No
Hsap\PTPN21
1 of 14
No
No
Hsap\RDX
1 of 14
No
No
1  
Hsap\TLN1
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (16)
Rnor\Frmd6
5 of 14
Yes
Yes
Rnor\Epb41l1
1 of 14
No
No
Rnor\Epb41l3
1 of 14
No
No
Rnor\Ezr
1 of 14
No
No
Rnor\Frmd1
1 of 14
No
Yes
Rnor\Frmd4b
1 of 14
No
No
Rnor\Frmd5
1 of 14
No
No
Rnor\Frmpd2
1 of 14
No
Yes
Rnor\Msn
1 of 14
No
No
Rnor\Nf2
1 of 14
No
No
Rnor\Ptpn3
1 of 14
No
No
Rnor\Ptpn4
1 of 14
No
No
Rnor\Ptpn13
1 of 14
No
No
Rnor\Ptpn14
1 of 14
No
No
Rnor\Ptpn21
1 of 14
No
No
Rnor\Rdx
1 of 14
No
No
Mus musculus (laboratory mouse) (18)
Mmus\Frmd6
4 of 14
Yes
Yes
Mmus\Epb41l1
1 of 14
No
No
Mmus\Epb41l3
1 of 14
No
No
Mmus\Epb41l4a
1 of 14
No
No
Mmus\Ezr
1 of 14
No
No
Mmus\Frmd3
1 of 14
No
No
Mmus\Frmd4a
1 of 14
No
No
Mmus\Frmd4b
1 of 14
No
No
Mmus\Frmd5
1 of 14
No
No
Mmus\Frmpd2
1 of 14
No
Yes
Mmus\Msn
1 of 14
No
No
Mmus\Nf2
1 of 14
No
No
Mmus\Ptpn3
1 of 14
No
No
Mmus\Ptpn4
1 of 14
No
No
Mmus\Ptpn13
1 of 14
No
No
Mmus\Ptpn14
1 of 14
No
No
Mmus\Ptpn21
1 of 14
No
No
Mmus\Rdx
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (11)
Xtro\frmd6
4 of 13
Yes
Yes
Xtro\epb41l3
1 of 13
No
No
Xtro\farp1
1 of 13
No
No
Xtro\frmd3
1 of 13
No
No
Xtro\frmd4a
1 of 13
No
Yes
Xtro\frmd4b
1 of 13
No
Yes
Xtro\frmd7
1 of 13
No
No
Xtro\LOC101731001
1 of 13
No
No
Xtro\mylip
1 of 13
No
No
Xtro\ptpn14
1 of 13
No
No
Xtro\ptpn21
1 of 13
No
No
Danio rerio (Zebrafish) (21)
Drer\zgc:172136
6 of 14
Yes
Yes
Drer\frmd6
4 of 14
No
Yes
Drer\epb41a
1 of 14
No
No
Drer\ezra
1 of 14
No
No
Drer\ezrb
1 of 14
No
No
Drer\frmd3
1 of 14
No
No
Drer\frmd4a
1 of 14
No
No
Drer\frmd4ba
1 of 14
No
No
Drer\frmd5b
1 of 14
No
No
Drer\frmpd2
1 of 14
No
No
Drer\LOC570903
1 of 14
No
No
Drer\msna
1 of 14
No
No
Drer\msnb
1 of 14
No
No
Drer\mylipa
1 of 14
No
No
Drer\nf2a
1 of 14
No
No
Drer\nf2b
1 of 14
No
No
Drer\ptpn3
1 of 14
No
No
Drer\ptpn4b
1 of 14
No
No
Drer\ptpn13
1 of 14
No
Yes
Drer\ptpn21
1 of 14
No
No
Drer\si:dkey-178k16.1
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (12)
Cele\frm-7
9 of 14
Yes
Yes
Cele\frm-5.2
2 of 14
No
Yes
Cele\epi-1
1 of 14
No
No
Cele\erm-1
1 of 14
No
No
Cele\F07C6.4
1 of 14
No
Yes
Cele\frm-1
1 of 14
No
No
Cele\frm-2
1 of 14
No
No
Cele\frm-3
1 of 14
No
No
Cele\frm-4
1 of 14
No
No
Cele\frm-5.1
1 of 14
No
Yes
Cele\nfm-1
1 of 14
No
No
Cele\ptp-1
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (12)
Agam\AgaP_AGAP009126
12 of 12
Yes
Yes
Agam\AgaP_AGAP001287
1 of 12
No
No
Agam\AgaP_AGAP001632
1 of 12
No
No
Agam\AgaP_AGAP004136
1 of 12
No
No
Agam\AgaP_AGAP007130
1 of 12
No
No
Agam\AgaP_AGAP007474
1 of 12
No
No
Agam\AgaP_AGAP008811
1 of 12
No
No
Agam\AgaP_AGAP009975
1 of 12
No
No
Agam\AgaP_AGAP010346
1 of 12
No
No
Agam\AgaP_AGAP012185
1 of 12
No
No
Agam\AgaP_AGAP013042
1 of 12
No
No
Agam\MOEH_ANOGA
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:ex. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (10)
Cdep
1 of 13
CG34347
1 of 13
cora
1 of 13
Frmd5
1 of 13
Mer
1 of 13
Moe
1 of 13
Pez
1 of 13
Ptpmeg
1 of 13
rhea
1 of 13
yrt
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 2 )
Allele
Disease
Evidence
References
exe1
model of  cancer
CEA with Mer4
(Hamaratoglu et al., 2006)
exAP49
model of  cancer
CEA
(Hamaratoglu et al., 2006)
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 6 )
Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
Homo sapiens (Human)
Gene name
Score
OMIM
OMIM Phenotype
DO term
Complementation?
Transgene?
FRMD1; FERM domain containing 1
7 of 14
      FRMD6; FERM domain containing 6
      4 of 14
      614555
          Functional Complementation Data
          Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
          Interactions
          Summary of Physical Interactions
          Interaction Browsers
          View in FlyBase Interactions Browser   View in MIST tool (external link)
          Please see the Physical Interaction reports below for full details
          RNA-RNA
          Physical Interaction
          Assay
          References
          ex - mir-278
          immunohistochemistry, luminiscence technology, necessary binding region
          (Easow et al., 2007, Teleman and Cohen, 2006)
          protein-protein
          Physical Interaction
          Assay
          References
          ex
          anti tag coimmunoprecipitation, anti tag western blot, far western blotting, tag visualisation, pull down, Identification by mass spectrometry
          (Wang et al., 2022, McCartney et al., 2000)
          ex - AP-2α
          anti tag coimmunoprecipitation, anti tag western blot
          (Liu et al., 2022)
          ex - Ack
          anti tag coimmunoprecipitation, anti tag western blot
          (Hu et al., 2016)
          ex - Cul1
          anti tag coimmunoprecipitation, peptide massfingerprinting
          (Ribeiro et al., 2014)
          ex - Dlish
          pull down, anti tag western blot, anti tag coimmunoprecipitation, Identification by mass spectrometry
          (Wang et al., 2019)
          ex - Hsp60A
          anti tag coimmunoprecipitation, peptide massfingerprinting
          (Kwon et al., 2013)
          ex - Lon
          anti tag coimmunoprecipitation, peptide massfingerprinting
          (Kwon et al., 2013)
          ex - Mec2
          anti tag coimmunoprecipitation, peptide massfingerprinting
          (Kwon et al., 2013)
          ex - Mer
          anti tag coimmunoprecipitation, western blot, anti tag western blot, far western blotting, tag visualisation
          (Li et al., 2024, Yang and Choi, 2021, McCartney et al., 2000)
          ex - POSH
          anti tag coimmunoprecipitation, anti tag western blot
          (Ma et al., 2018)
          ex - Patronin
          anti tag coimmunoprecipitation, western blot
          (Yang and Choi, 2021)
          ex - Ptp61F
          anti tag coimmunoprecipitation, anti tag western blot
          (Liu et al., 2022)
          ex - Ran
          anti tag coimmunoprecipitation, peptide massfingerprinting
          (Kwon et al., 2013)
          ex - Schip1
          anti bait coimmunoprecipitation, anti tag western blot, pull down
          (Chung et al., 2016)
          ex - crb
          pull down, anti tag western blot, anti tag coimmunoprecipitation
          (Ribeiro et al., 2014, Ling et al., 2010)
          ex - drk
          anti tag coimmunoprecipitation, peptide massfingerprinting
          (Kwon et al., 2013)
          ex - ed
          anti tag coimmunoprecipitation, anti tag western blot
          (Yue et al., 2012)
          ex - ft
          anti tag coimmunoprecipitation, western blot, proximity ligation assay, fluorescence microscopy, pull down, anti tag western blot
          (Fulford et al., 2023)
          ex - gish
          anti tag coimmunoprecipitation, anti tag western blot, Identification by mass spectrometry
          (Fulford et al., 2019)
          ex - hpo
          two hybrid, anti tag coimmunoprecipitation, anti tag western blot
          (Reddy and Irvine, 2011, Yu et al., 2010)
          ex - kibra
          anti tag coimmunoprecipitation, anti tag western blot, anti bait coimmunoprecipitation, western blot
          (Genevet et al., 2010, Yu et al., 2010)
          ex - kst
          anti tag coimmunoprecipitation, anti tag western blot
          (Fletcher et al., 2015)
          ex - nonA
          anti tag coimmunoprecipitation, peptide massfingerprinting
          (Kwon et al., 2013)
          ex - sav
          anti tag coimmunoprecipitation, anti tag western blot
          (Yu et al., 2010)
          ex - slmb
          anti tag coimmunoprecipitation, peptide massfingerprinting, pull down, anti tag western blot, western blot
          (Fulford et al., 2019, Wang et al., 2019, Zhang et al., 2015, Ribeiro et al., 2014)
          ex - wts
          anti tag coimmunoprecipitation, anti tag western blot, western blot
          (Sun et al., 2015)
          ex - yki
          anti tag coimmunoprecipitation, western blot, anti bait coimmunoprecipitation, anti tag western blot, Identification by mass spectrometry, pull down, peptide massfingerprinting
          (Wang et al., 2022, Srivastava et al., 2021, Hu et al., 2016, Kwon et al., 2013, Badouel et al., 2009, Oh et al., 2009)
          Summary of Genetic Interactions
          Interaction Browsers
          View in FlyBase Interactions Browser   View in MIST tool (external link)
          Please look at the allele data for full details of the genetic interactions
          Starting gene(s)
          Interaction type
          Interacting gene(s)
          Reference
          ex
          suppressible
          crb
          (Fulford et al., 2019)
          ex
          suppressible
          CycD|Cdk4
          (Hipfner et al., 2002)
          ex
          suppressible
          hpo
          (Hamaratoglu et al., 2006)
          ex
          suppressible
          jub
          (Rauskolb et al., 2011)
          ex
          enhanceable
          kibra
          (Lucas et al., 2013, Yu et al., 2010)
          ex
          suppressible
          kibra
          (Tokamov et al., 2021, Su et al., 2017, Genevet et al., 2010)
          ex
          enhanceable
          Mer
          (McCartney et al., 2000)
          ex
          suppressible
          mir-278
          (Teleman and Cohen, 2006)
          ex
          suppressible
          mir-ban
          (Hipfner et al., 2002)
          ex
          enhanceable
          Mnat9
          (Mok and Choi, 2022)
          ex
          suppressible
          Mnat9
          (Mok and Choi, 2022)
          ex
          enhanceable
          mop
          (Gilbert et al., 2011)
          ex
          enhanceable
          Myc
          (Ziosi et al., 2010)
          ex
          suppressible
          par-1
          (Huang et al., 2013)
          ex|mtm
          suppressible
          Pi3K68D
          (Hu et al., 2022)
          ex|mtm
          suppressible
          Rab11
          (Hu et al., 2022)
          ex|mtm
          suppressible
          Rok
          (Hu et al., 2022)
          ex
          suppressible
          Schip1
          (Chung et al., 2016)
          ex
          suppressible
          sd
          (Koontz et al., 2013)
          ex
          suppressible
          tai
          (Zhang et al., 2015)
          ex
          suppressible
          Tao
          (Poon et al., 2011)
          ex
          suppressible
          Tgi
          (Koontz et al., 2013)
          ex
          suppressible
          wg
          (Pellock et al., 2007)
          ex
          suppressible
          wts
          (Cho et al., 2006, Hamaratoglu et al., 2006)
          ex
          enhanceable
          wts
          (Hamaratoglu et al., 2006)
          ex
          suppressible
          Zyx
          (Gaspar et al., 2015, Rauskolb et al., 2011)
          Starting gene(s)
          Interaction type
          Interacting gene(s)
          Reference
          Apc2|Apc
          suppressible
          ex
          (Suijkerbuijk et al., 2016)
          crb
          suppressible
          ex
          (Su et al., 2017, Chung et al., 2016)
          CycE
          suppressible
          ex
          (Brumby et al., 2004)
          Dcr-2|Snr1
          suppressible
          ex
          (Jeibmann et al., 2014)
          dia
          suppressible
          ex
          (Sansores-Garcia et al., 2011)
          Dlish
          suppressible
          ex
          (Misra and Irvine, 2016)
          dsh
          enhanceable
          ex
          (Blaumueller and Mlodzik, 2000)
          ft
          enhanceable
          ex
          (Silva, 2006)
          ft
          suppressible
          ex
          (Willecke et al., 2006)
          kibra
          enhanceable
          ex
          (Lucas et al., 2013, Baumgartner et al., 2010, Yu et al., 2010)
          kibra
          suppressible
          ex
          (Su et al., 2017, Genevet et al., 2010)
          Kr
          enhanceable
          ex
          (Abrell et al., 2000)
          l(3)mbt
          suppressible
          ex
          (Richter et al., 2011)
          Mer
          enhanceable
          ex
          (Willecke et al., 2006, McCartney et al., 2000)
          Mnat9
          enhanceable
          ex
          (Mok and Choi, 2022)
          Mnat9
          suppressible
          ex
          (Mok and Choi, 2022)
          mop
          suppressible
          ex
          (Gilbert et al., 2011)
          mop
          enhanceable
          ex
          (Gilbert et al., 2011)
          mtm
          enhanceable
          ex
          (Hu et al., 2022)
          obk
          suppressible
          ex
          (Atallah et al., 2004)
          Patronin
          suppressible
          ex
          (Yang and Choi, 2021)
          pyd
          enhanceable
          ex
          (Djiane et al., 2011)
          Snr1
          enhanceable
          ex
          (Marenda et al., 2004)
          Snr1
          suppressible
          ex
          (Jeibmann et al., 2014)
          TER94
          suppressible
          ex
          (Li et al., 2024)
          wts
          enhanceable
          ex
          (Hamaratoglu et al., 2006)
          wts
          suppressible
          ex
          (Badouel et al., 2009)
          yki
          suppressible
          ex|hpo
          (Oh et al., 2009)
          yki
          suppressible
          ex|wts|hpo
          (Oh et al., 2009)
          External Data
          Subunit Structure (UniProtKB)
          Forms a complex with Kibra and Mer (PubMed:20159598). Interacts (via RXPPXY motif) with Kibra (via domain WW 1) (PubMed:20159598). Interacts with Mer and Hpo (via SARAH domain) (PubMed:20159598). Interacts with Schip1; the interaction results in recruitment of Schip1 to the apical cell membrane (PubMed:26954546). Interacts with ack and yki (PubMed:27462444).
          (UniProt, Q07436 )
          Linkouts
          BioGRID - A database of protein and genetic interactions.
          DroID - A comprehensive database of gene and protein interactions.
          MIST (genetic) - An integrated Molecular Interaction Database
          MIST (protein-protein) - An integrated Molecular Interaction Database
          Pathways
          Signaling Pathways (FlyBase)
          Metabolic Pathways
          FlyBase
          External Links
          External Data
          Linkouts
          KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
          Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
          Class of Gene
          Genomic Location and Detailed Mapping Data
          Chromosome (arm)
          2L
          Recombination map
          2-0.7
          Cytogenetic map
          21C2-21C2
          Sequence location
          FlyBase Computed Cytological Location
          Cytogenetic map
          Evidence for location
          21C2-21C2
          Limits computationally determined from genome sequence between P{lacW}l(2)k09610k09610&P{lacW}Sk09538a and P{PZ}l(2)1068510685&P{lacW}Tango14k00619
          Experimentally Determined Cytological Location
          Cytogenetic map
          Notes
          References
          21C4-21C5
          (determined by in situ hybridisation)
          (Spradling et al., 1999)
          21C-21C
          (determined by in situ hybridisation)
          (Kania et al., 1995)
          21C3-21C4
          21C4--5
          (BDGP Project Members, 1994-1999)
          21C1-21C4
          (determined by in situ hybridisation)
          (Boedigheimer and Laughon, 1993)
          Experimentally Determined Recombination Data
          Location

          2-0.7

          (FlyBase, 2016)

          2-0.0

          (Comeron et al., 2012)

          2-0.65

          (Huettner and Bryant, 1988)

          2-0.1

          Left of (cM)
          Right of (cM)
          Notes

          Mapped with exl2ey.

          (Huettner and Bryant, 1988)
          Stocks and Reagents
          Stocks (35)
          Genomic Clones (16)
           

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

          cDNA Clones (19)
           

          Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see JBrowse for alignment of the cDNAs and ESTs to the gene model.

          cDNA clones, fully sequenced
          BDGP DGC clones
          Other clones
            Drosophila Genomics Resource Center cDNA clones

            For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

            cDNA Clones, End Sequenced (ESTs)
            BDGP DGC clones
            Other clones
            RNAi and Array Information
            Linkouts
            DRSC - Results frm RNAi screens
            Antibody Information
            Laboratory Generated Antibodies
             

            polyclonal

            (Hu et al., 2016, Maitra et al., 2006, Boedigheimer and Laughon, 1993)
            Commercially Available Antibodies
             
            Cell Line Information
            Publicly Available Cell Lines
             
              Other Stable Cell Lines
               
                Other Comments

                dsRNA made from templates generated with primers directed against this gene is tested in an S2 cell phosphorylation experiment to assess teh hpo signal transduction pathway.

                (Willecke et al., 2006)

                The actions of ex are epistatic to ft.

                (Willecke et al., 2006)

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

                (Kiger et al., 2003)

                ex has a common role in all disc tissues and can negatively regulate tissue growth, induce apoptosis, and perturb planar polarity.

                (Blaumueller and Mlodzik, 2000)

                Mer and ex function cooperatively in regulating cell proliferation and differentiation in developing epithelial cells.

                (McCartney et al., 2000)

                Identification: Enhancer trap expression pattern survey for loci expressed in the ring gland.

                (Harvie et al., 1998)

                Clonal analysis reveals that ex functions in the apical cell domain to regulate the growth rate of imaginal discs.

                (Boedigheimer et al., 1997)

                Variation of a microsatellite within the ex locus has been studied in North American populations of D.melanogaster.

                (Goldstein and Clark, 1995)

                Mutants display hyperplastic phenotype, with imaginal disc overgrowth.

                (Mechler, 1994)

                The ex gene is necessary for proper growth control of imaginal discs.

                (Boedigheimer and Laughon, 1993)

                Presumed additional alleles obtained by mobilization of P-element. All are recessive lethals that die during early larval stages.

                Relationship to Other Genes
                Source for database merge of

                Source for merge of: ex l(2)01270

                (Harvie et al., 1998)

                Source for merge of: ex l(2)k12913

                (Spradling et al., 1999)

                Source for merge of: ex brl

                (Scott, 1995.4.5)
                Additional comments
                Nomenclature History
                Source for database identify of

                Source for identity of: ex CG4114

                (Guan, 1999.11)
                Nomenclature comments
                Etymology
                Synonyms and Secondary IDs (13)
                Reported As
                Symbol Synonym
                1270
                (Harvie et al., 1998)
                CG4114
                (National Institute of Genetics Fly Stocks, 2022-, Trivedi et al., 2020, Balakrishnan et al., 2015, Hosono et al., 2015, Jeibmann et al., 2014, Ni et al., 2011.7.19, Ida et al., 2009, Keleman et al., 2009.8.5, Perkins et al., 2009, Molnar et al., 2006, Metaxakis et al., 2005)
                EX
                (Grifoni et al., 2013, Harvey and Hariharan, 2012)
                Ex
                (Bonello et al., 2023, Petsakou et al., 2023, Kong et al., 2022, Liu et al., 2022, Cho and Jiang, 2021, Gogia et al., 2021, Johnson and Leatherman, 2021, Fulford and McNeill, 2020, Gou et al., 2020, Mohammadi et al., 2020, Fulford et al., 2019, Gao et al., 2019, Gokhale and Pfleger, 2019, Khadilkar and Tanentzapf, 2019, Bae and Luo, 2018, Elbediwy and Thompson, 2018, Fulford et al., 2018, Baker, 2017, Kulaberoglu et al., 2017, Richardson and Portela, 2017, Fallahi et al., 2016, Gunn-Moore et al., 2016, Hu et al., 2016, Jagannathan et al., 2016, Yadav et al., 2016, Bae et al., 2015, Cao et al., 2015, Flores-Benitez and Knust, 2015, Gaspar et al., 2015, Irvine and Harvey, 2015, Thompson and Sahai, 2015, Wang and Baker, 2015, Zheng et al., 2015, Hu et al., 2014, Ribeiro et al., 2014, Sadeqzadeh et al., 2014, Sing et al., 2014, Sytnikova et al., 2014, Andersen et al., 2013, Guo et al., 2013, Huang et al., 2013, Jukam et al., 2013, Khan et al., 2013, Kwon et al., 2013, Lucas et al., 2013, Matsui and Lai, 2013, Wehr et al., 2013, Yu and Guan, 2013, Angus et al., 2012, Fausti et al., 2012, Jin et al., 2012, Kagey et al., 2012, Liu et al., 2012, Poernbacher et al., 2012, Yue et al., 2012, Boggiano et al., 2011, Chan et al., 2011, Genevet and Tapon, 2011, Gilbert et al., 2011, Halder and Johnson, 2011, Laprise, 2011, Salah and Aqeilan, 2011, Zhao et al., 2011, Ren et al., 2010, Badouel et al., 2009, Hamaratoglu et al., 2009, Zhang et al., 2009, Wu et al., 2008, Hariharan, 2006, Polesello et al., 2006)
                Expanded
                (Levayer and Moreno, 2013, Fausti et al., 2012, Gilbert et al., 2011, Laprise, 2011, Salah and Aqeilan, 2011)
                Frmd6
                (van Soldt and Cardoso, 2020)
                brl
                ex
                (Yu et al., 2025, Guo et al., 2024, Kroeger et al., 2024, Li et al., 2024, Luo et al., 2024, Portela et al., 2024, Sun et al., 2024, Chan et al., 2023, Kim et al., 2023, Yamada et al., 2023, Gao et al., 2022, Hu et al., 2022, Kumar and Baker, 2022, Li et al., 2022, Perlegos et al., 2022, Wang et al., 2022, Yang et al., 2022, Ahmad et al., 2021, Badmos et al., 2021, Ding et al., 2021, Ferguson et al., 2021, Gong et al., 2021, Srivastava et al., 2021, Tokamov et al., 2021, Xu et al., 2021, Yang and Choi, 2021, Aguilar-Aragon et al., 2020, Baker, 2020, Deng et al., 2020, Earl et al., 2020, Guo et al., 2020, Mugat et al., 2020, Skouloudaki et al., 2020, Sun et al., 2020, Wang and Spradling, 2020, Yu et al., 2020, Alégot et al., 2019, Chang et al., 2019, Li et al., 2019, Liu et al., 2019, Ma et al., 2019, Misra and Irvine, 2019, Nan et al., 2019, Sarpal et al., 2019, Snigdha et al., 2019, Wang and Baker, 2019, Wang et al., 2019, Wong et al., 2019, Xu et al., 2019, Yee et al., 2019, Zhang et al., 2019, Albert et al., 2018, Cho et al., 2018, Fletcher et al., 2018, Katsukawa et al., 2018, Lee et al., 2018, Tsai et al., 2018, Wang and Baker, 2018, Wang et al., 2018, Yu and Pan, 2018, Anderson et al., 2017, Hevia et al., 2017, Martín et al., 2017, Pascual et al., 2017, Shu and Deng, 2017, Su et al., 2017, Yamamoto et al., 2017, Zhang et al., 2017, Zheng et al., 2017, Boone et al., 2016, Chung et al., 2016, Deng et al., 2016, Gene Disruption Project members, 2016-, Huang et al., 2016, Kockel et al., 2016, Kwan et al., 2016, Misra and Irvine, 2016, Pan et al., 2016, Shih et al., 2016, Suijkerbuijk et al., 2016, Bosch et al., 2015, Dent et al., 2015, Doggett et al., 2015, Dong et al., 2015, Enomoto et al., 2015, Fletcher et al., 2015, Hirabayashi and Cagan, 2015, Jia et al., 2015, Keder et al., 2015, Kwon et al., 2015, Li et al., 2015, Meng et al., 2015, Meserve and Duronio, 2015, Parker and Struhl, 2015, Su, 2015, Sun et al., 2015, Vrabioiu and Struhl, 2015, Zaessinger et al., 2015, Zhang et al., 2015, Zhang et al., 2015, Zhu et al., 2015, Ashwal-Fluss et al., 2014, Bossuyt et al., 2014, Huang and Kalderon, 2014, Huang et al., 2014, Qing et al., 2014, Rauskolb et al., 2014, Robbins et al., 2014, Tipping and Perrimon, 2014, Christiansen et al., 2013, Degoutin et al., 2013, Deng et al., 2013, Huang et al., 2013, Ilanges et al., 2013, Jukam et al., 2013, Khan et al., 2013, Koontz et al., 2013, Kwon et al., 2013, Losick et al., 2013, Lucas et al., 2013, Marcinkevicius and Zallen, 2013, McKay and Lieb, 2013, Oh et al., 2013, Ohtani et al., 2013, Perea et al., 2013, Sidor et al., 2013, Webber et al., 2013, Yu et al., 2013, Chen and Verheyen, 2012, Chen et al., 2012, Fletcher et al., 2012, Hafezi et al., 2012, Herranz et al., 2012, Matakatsu and Blair, 2012, Zhang et al., 2012, Bao et al., 2011, Djiane et al., 2011, Fernández et al., 2011, Friedman et al., 2011, Gilbert et al., 2011, Grusche et al., 2011, Jukam and Desplan, 2011, Laprise, 2011, Poon et al., 2011, Rauskolb et al., 2011, Reddy and Irvine, 2011, Richter et al., 2011, Slattery et al., 2011, Sun et al., 2011, Yan et al., 2011, Zhang et al., 2011, Baumgartner et al., 2010, Baumgartner et al., 2010, Chen et al., 2010, Genevet et al., 2010, Genevet et al., 2010, Grzeschik et al., 2010, Kühnlein, 2010, Ling et al., 2010, Milton et al., 2010, Morrison and Halder, 2010, Neto-Silva et al., 2010, Reddy et al., 2010, Ribeiro et al., 2010, Robinson et al., 2010, Shaw et al., 2010, Terriente-Félix et al., 2010, Yu et al., 2010, Yu et al., 2010, Zecca and Struhl, 2010, Ziosi et al., 2010, Classen et al., 2009, Genevet et al., 2009, Ida et al., 2009, Oh and Irvine, 2009, Sopko et al., 2009, Zhang et al., 2009, Lee et al., 2008, Oh and Irvine, 2008, Rogulja et al., 2008, Sun et al., 2008, Willecke et al., 2008, Yu et al., 2008, Zhang et al., 2008, Zhang et al., 2008, Zhao et al., 2008, Acevedo et al., 2007, Feng and Irvine, 2007, Feng et al., 2007, Meignin et al., 2007, Pellock et al., 2007, Polesello and Tapon, 2007, Tyler and Baker, 2007, Tyler et al., 2007, Zhao et al., 2007, Bennett and Harvey, 2006, Cho et al., 2006, Hamaratoglu et al., 2006, Hamaratoglu et al., 2006, Maitra et al., 2006, Maitra et al., 2006, Molnar et al., 2006, Polesello et al., 2006, Silva, 2006, Willecke et al., 2006, McCartney et al., 2000)
                exp
                (Bajpai et al., 2020)
                expanded
                (Fletcher et al., 2015, Ilanges et al., 2013)
                l(2)01270
                (Abrell et al., 2000)
                l(2)ey
                (Huettner and Bryant, 1988)
                Name Synonyms
                Expanded
                (Gou et al., 2023, Aguilar-Aragon et al., 2020, van Soldt and Cardoso, 2020, Romine et al., 2019, Chan et al., 2016, Gunn-Moore et al., 2016, Misra and Irvine, 2016, Suijkerbuijk et al., 2016, Zheng et al., 2015, Bossuyt et al., 2014, Sadeqzadeh et al., 2014, Sing et al., 2014, Sorrentino et al., 2014, Degoutin et al., 2013, Deng et al., 2013, Harvey et al., 2013, Khan et al., 2013, Konsavage and Yochum, 2013, Lucas et al., 2013, Yu and Guan, 2013, Tepass, 2012, Bao et al., 2011, Boggiano et al., 2011, Halder and Johnson, 2011, Reddy and Irvine, 2011, Baumgartner et al., 2010, Genevet et al., 2010, Grzeschik et al., 2010, Ling et al., 2010, Reddy et al., 2010, Robinson et al., 2010, Yu et al., 2010, Badouel et al., 2009, Hamaratoglu et al., 2009, Mao et al., 2009, Oh et al., 2009, Badouel et al., 2008, Oh and Irvine, 2008, Harvey and Bennett, 2007, Polesello and Tapon, 2007, Willecke et al., 2007, Herz et al., 2006, Polesello et al., 2006, Silva et al., 2006, Thompson and Cohen, 2006, Willecke et al., 2006, Sano et al., 2005)
                braille
                expanded
                (Buckley and St Johnston, 2022, Mugat et al., 2020, Khadilkar and Tanentzapf, 2019, Li et al., 2019, Hevia et al., 2017, Jonchère et al., 2017, Kwon et al., 2015, Meng et al., 2015, Wang and Baker, 2015, Ferguson and Martinez-Agosto, 2014, Hu et al., 2014, Ikmi et al., 2014, Jeibmann et al., 2014, Robbins et al., 2014, Christiansen et al., 2013, Enderle and McNeill, 2013, Huang et al., 2013, Jukam et al., 2013, Lawrence and Casal, 2013, Losick et al., 2013, Lucas et al., 2013, McKay and Lieb, 2013, Chen et al., 2012, Sienski et al., 2012, Zhang et al., 2012, Grusche et al., 2011, Rauskolb et al., 2011, Slattery et al., 2011, Chen et al., 2010, Kühnlein, 2010, Milton et al., 2010, Morrison and Halder, 2010, Neto-Silva et al., 2010, Terriente-Félix et al., 2010, Zecca and Struhl, 2010, Jukam and Claude, 2008, Sun et al., 2008, Yu et al., 2008, Feng and Irvine, 2007, Hamaratoglu et al., 2007, Jukam and Desplan, 2007, Meignin et al., 2007, Pellock et al., 2007, Tyler and Baker, 2007, Tyler et al., 2007, Cho et al., 2006, Edgar, 2006, Hariharan, 2006, Maitra et al., 2006, McCartney et al., 2000, Edwards et al., 1997, Boedigheimer, 1996.12.17)
                Secondary FlyBase IDs
                • FBgn0000220
                • FBgn0000610
                • FBgn0002118
                • FBgn0010479
                • FBgn0021897
                Datasets (1)
                Study focus (1)
                Experimental Role
                Project
                Project Type
                Title
                • bait_protein
                Hippo_Pathway_interaction_map
                Interaction map generated by purification of Hippo pathway factors, with identification of copurifying proteins by mass spectrometry.
                Study result (0)
                Result
                Result Type
                Title
                External Crossreferences and Linkouts ( 54 )
                Sequence Crossreferences
                NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
                GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
                GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
                RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
                UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
                UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
                UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
                Other crossreferences
                AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
                DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
                EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
                FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
                FlyMine - An integrated database for Drosophila genomics
                InterPro - A database of protein families, domains and functional sites
                KEGG Genes - Molecular building blocks of life in the genomic space.
                MARRVEL_MODEL - MARRVEL (model organism gene)
                Linkouts
                BioGRID - A database of protein and genetic interactions.
                Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
                DroID - A comprehensive database of gene and protein interactions.
                DRSC - Results frm RNAi screens
                Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
                FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
                FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
                Flygut - An atlas of the Drosophila adult midgut
                iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
                Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
                MIST (genetic) - An integrated Molecular Interaction Database
                MIST (protein-protein) - An integrated Molecular Interaction Database
                Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                References (472)