FB2025_05 , released December 11, 2025
Gene: Dmel\dally
Open Close
General Information
Symbol
Dmel\dally
Species
D. melanogaster
Name
division abnormally delayed
Annotation Symbol
CG4974
Feature Type
protein_coding_gene
FlyBase ID
FBgn0263930
Gene Model Status
Current
Stock Availability
126 publicly available
Gene Summary
division abnormally delayed (dally) encodes a core protein of heparan sulfate proteoglycans of the glypican family. It acts as a co-receptor for growth factors and morphogens, such as the products of dpp, wg, hh, upd1 and FGFs, affecting signaling and distribution of these ligands. The roles of the product of dally include wing development and germline stem cell maintenance. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Also Known As

gem, l(3)SG18

Key Links
Genomic Location
Cytogenetic map
66E1-66E3
Sequence location
Recombination map
3-27
RefSeq locus
NT_037436 REGION:8827505..8891192
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 (39 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables protein binding
inferred from physical interaction with FLYBASE:nord; FB:FBgn0050418
(Akiyama et al., 2022)
inferred from physical interaction with FLYBASE:dpp; FB:FBgn0000490
(Kirkpatrick et al., 2006)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (31 terms)
Terms Based on Experimental Evidence (28 terms)
CV Term
Evidence
References
involved_in axonogenesis
inferred from mutant phenotype
(Grueber et al., 2007)
involved_in chaeta development
inferred from mutant phenotype
(Kirkpatrick et al., 2006)
involved_in chaeta morphogenesis
inferred from mutant phenotype
(Takeo et al., 2005)
involved_in compound eye development
inferred from mutant phenotype
(Kirkpatrick et al., 2006)
involved_in dendrite morphogenesis
inferred from mutant phenotype
(Grueber et al., 2007)
involved_in epithelial cell migration, open tracheal system
inferred from genetic interaction with FLYBASE:dlp; FB:FBgn0041604
(Yan and Lin, 2007)
involved_in establishment of protein localization to extracellular region
inferred from mutant phenotype
(Simon et al., 2024)
involved_in female germ-line stem cell asymmetric division
inferred from mutant phenotype
(Guo and Wang, 2009)
involved_in germ-line stem cell population maintenance
inferred from mutant phenotype
(Guo and Wang, 2009)
involved_in imaginal disc-derived leg morphogenesis
inferred from mutant phenotype
(Takeo et al., 2005)
involved_in imaginal disc-derived wing vein morphogenesis
inferred from mutant phenotype
(Kirkpatrick et al., 2006, Takeo et al., 2005)
involved_in maintenance of protein location in extracellular region
inferred from mutant phenotype
(Bilioni et al., 2013, Ayers et al., 2012, Ayers et al., 2010, Fujise et al., 2003)
involved_in motor neuron axon guidance
inferred from mutant phenotype
(Cho et al., 2012)
involved_in negative regulation of receptor internalization
inferred from mutant phenotype
(Simon et al., 2024)
involved_in negative regulation of semaphorin-plexin signaling pathway
inferred from genetic interaction with FLYBASE:Sema1a; FB:FBgn0011259
(Cho et al., 2012)
involved_in positive regulation of BMP signaling pathway
inferred from mutant phenotype
(Ell et al., 2024, Simon et al., 2024, Akiyama et al., 2022, Dejima et al., 2011, Guo and Wang, 2009, Crickmore and Mann, 2007, Takeo et al., 2005, Fujise et al., 2003)
involved_in positive regulation of canonical Wnt signaling pathway
inferred from genetic interaction with FLYBASE:Notum; FB:FBgn0044028
(Han et al., 2005)
involved_in positive regulation of fibroblast growth factor receptor signaling pathway
inferred from genetic interaction with FLYBASE:dlp; FB:FBgn0041604
(Yan and Lin, 2007)
involved_in positive regulation of smoothened signaling pathway
inferred from genetic interaction with FLYBASE:dlp; FB:FBgn0041604
(Avanesov and Blair, 2013)
inferred from mutant phenotype
(Bilioni et al., 2013, Ayers et al., 2012, Ayers et al., 2010, Takeo et al., 2005)
involved_in regulation of cell cycle
inferred from mutant phenotype
(Nakato et al., 2002)
involved_in regulation of imaginal disc growth
inferred from mutant phenotype
(Crickmore and Mann, 2007, Hufnagel et al., 2007)
involved_in segment polarity determination
inferred from mutant phenotype
(Baeg et al., 2001)
involved_in sensory organ boundary specification
inferred from mutant phenotype
(Fujise et al., 2001)
involved_in wing disc anterior/posterior pattern formation
inferred from mutant phenotype
(Fujise et al., 2003)
involved_in wing disc development
inferred from mutant phenotype
(Kirkpatrick et al., 2006)
involved_in wing disc dorsal/ventral pattern formation
inferred from mutant phenotype
(Han et al., 2005)
inferred from genetic interaction with FLYBASE:wg; FB:FBgn0284084
(Han et al., 2005)
involved_in wing disc morphogenesis
inferred from mutant phenotype
(Takeo et al., 2005)
involved_in wing disc pattern formation
inferred from mutant phenotype
(Ayers et al., 2012, Ayers et al., 2010)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
involved_in cell migration
inferred from biological aspect of ancestor with PANTHER:PTN002247069
(GO Reference Genome Project, 2011-)
involved_in positive regulation of canonical Wnt signaling pathway
inferred from biological aspect of ancestor with PANTHER:PTN000850517
(GO Reference Genome Project, 2011-)
involved_in regulation of protein localization to membrane
inferred from biological aspect of ancestor with PANTHER:PTN002247069
(GO Reference Genome Project, 2011-)
involved_in regulation of signal transduction
inferred from electronic annotation with InterPro:IPR001863
(InterPro Project Members, 2004-)
Cellular Component (7 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
located_in apical plasma membrane
inferred from direct assay
(Ayers et al., 2010)
located_in basolateral plasma membrane
inferred from direct assay
(Ayers et al., 2010)
located_in cell surface
inferred from direct assay
(Kirkpatrick et al., 2006)
is_active_in external side of plasma membrane
inferred from direct assay
(Ell et al., 2024)
located_in external side of plasma membrane
inferred from mutant phenotype
(Ayers et al., 2012)
located_in perivitelline space
inferred from high throughput direct assay
(Lye et al., 2014)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
is_active_in cell surface
inferred from biological aspect of ancestor with PANTHER:PTN002247069
(GO Reference Genome Project, 2011-)
located_in extracellular matrix
inferred from electronic annotation with InterPro:IPR001863
(InterPro Project Members, 2004-)
located_in plasma membrane
inferred from electronic annotation with InterPro:IPR001863
(InterPro Project Members, 2004-)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
Belongs to the glypican family. (Q24114)
Protein Signatures (InterPro)
Summaries
Gene Snapshot
division abnormally delayed (dally) encodes a core protein of heparan sulfate proteoglycans of the glypican family. It acts as a co-receptor for growth factors and morphogens, such as the products of dpp, wg, hh, upd1 and FGFs, affecting signaling and distribution of these ligands. The roles of the product of dally include wing development and germline stem cell maintenance. [Date last reviewed: 2019-03-07]
Pathway (FlyBase)
POSITIVE REGULATORS OF WNT-TCF SIGNALING PATHWAY -
Positive regulators of Wnt-TCF (canonical Wnt) signaling up-regulate the pathway, resulting in the enhancement of transcriptional regulation mediated by β-catenin (arm).
POSITIVE REGULATORS OF FGFR SIGNALING PATHWAY -
Positive regulators of Fibroblast Growth Factor Receptor signaling up-regulate the pathway, enhancing the activation of ERK kinase (rl) or acting on other effectors.
POSITIVE REGULATORS OF HEDGEHOG SIGNALING PATHWAY -
Positive regulators of hedgehog signaling up-regulate the pathway, resulting in the activation of transcription of hh-responsive genes.
POSITIVE REGULATORS OF BMP SIGNALING PATHWAY -
Positive regulators of Bone Morphogenetic Protein (BMP) signaling up-regulate the pathway, ultimately resulting in the increased nuclear activity of the Mad/Med transcription factor complex.
Gene Group (FlyBase)
GLYPICANS -
Glypicans are glycosylphosphatidylinositol (GPI) anchored heparan sulfate proteoglycans. Glypicans can bind growth factors and morphogens, regulating their extracellular distribution. (Adapted from FBrf0138451).
Protein Function (UniProtKB)
Cell surface proteoglycan that bears heparan sulfate (PubMed:35609633, PubMed:8582281). Functions as a coreceptor for growth factors and morphogens, such as the products of dpp, to regulate signaling and distribution of these ligands (PubMed:35609633, PubMed:8582281). Required for cell division patterning during postembryonic development of the nervous system (PubMed:8582281). Plays a role in dpp/BMP signaling possibly by stabilizing dpp and thereby creating a morphological gradient during wing development (PubMed:35609633). Might have a role in testis development (PubMed:35609633). Functions with magu and fwe in a mechanism of scaling, which utilises apoptosis to ensure that the dpp patterning gradient remains proportional to the size of the growing wing (PubMed:35301437). In this mechanism, fwe represses dally and Magu-dependent activity in expanding the gradient, and dally/Magu inhibits fwe-dependent apoptosis to keep cell death rate low (PubMed:35301437). When the levels of these different proteins are optimally regulated the gradient correctly scales with organ growth but when this fails, fwe-mediated apoptosis is activated to trim the developing tissue to match the correct size of the gradient (PubMed:35301437).
(UniProt, Q24114)
Summary (Interactive Fly)

membrane bound glypican related proteoglycan - effects cell division of lamina precursor cells of the optic lobe - Ras/MEK/MAPK-mediated regulation of heparin sulphate proteoglycans promotes retinal fate in the Drosophila eye-antennal disc - Dally mediates the autonomous and nonautonomous effects on tissue growth caused by activation of the PI3K and TOR pathways

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

Please see the JBrowse view of Dmel\dally 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 Q24114)

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

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.46

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
dally-RA
FBtr0076583
NM_079259
3444
626
dally-RB
FBtr0305901
NM_001259756
4483
626
Additional Transcript Data and Comments
Reported size (kB)

4.4 (northern blot)

(Nakato et al., 1995)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
dally-PA
FBpp0076310
69.0
626
6.39
Q24114
NP_523983
AAF50358
dally-PB
FBpp0297069
69.0
626
6.39
Q24114
NP_001246685
AFH04356
Polypeptides with Identical Sequences

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

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

626 (aa)

(Nakato et al., 1995)
Comments
External Data
Subunit Structure (UniProtKB)

Interacts with nord; the interaction promotes dally degradation (PubMed:35609633). Interacts with Magu (PubMed:35301437). As part of the dally/ Magu complex, associates with fwe (isoforms ubi, LoseA and LoseB) and is unable to interact with fwe independently of Magu (PubMed:35301437).

(UniProt, Q24114)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
UniProt
Mapped Features

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

External Data
Crossreferences
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).

-1.40

Transcript Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
oogenesis stage S4 -- S10
oocyte associated follicular epithelium
(Hayashi et al., 2012)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Nakato et al., 1995)
embryonic stage 2
organism | ubiquitous
(Lin and Perrimon, 1999)
embryonic stage 8
ectoderm | restricted
(Lin and Perrimon, 1999)
organism | segment polarity expression pattern
(Lin and Perrimon, 1999)
embryonic stage 9
embryonic epidermis | segment polarity expression pattern
(Tsuda et al., 1999)
third instar larval stage
lamina anlage
(Nakato et al., 1995)
larval inner optic anlage
(Nakato et al., 1995)
leg disc
(Nakato et al., 1995)
eye-antennal disc
(Nakato et al., 1995)
wing disc
(Nakato et al., 1995)
adult stage | female
germarium | posterior
(Hayashi et al., 2012)
stalk follicle cell | precursor
(Hayashi et al., 2012)
polar follicle cell | precursor
(Hayashi et al., 2012)
Additional Descriptive Data

dally transcript accumulates in somatic cells at the posterior of the germarium, including precursors to both interfollicle and polar follicle cells. dally is also expressed in the oocyte associated follicular epithelium.

(Hayashi et al., 2012)

dally transcripts are uniformly expressed at early embryonic stages. At stage 8, a segmentally repeated expression pattern is observed that occurs 3-4 cells anterior to the wg-expressing cells.

(Lin and Perrimon, 1999)

dally transcripts are found in the lamina precursor cells (LPCs) in third instar larval brains. The highest level occurs in LPCs along the anterior segment of the lamina furrow in cells that are in the G2 and M phase of the first division. Expression is also seen in the inner optic anlage and in some cells of the central brain. Expression also occurs in the antenna, eye, leg, and wing discs and in embryos.

(Nakato et al., 1995)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult head
(Aradska et al., 2015)
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in apical plasma membrane
inferred from direct assay
(Ayers et al., 2010)
located_in basolateral plasma membrane
inferred from direct assay
(Ayers et al., 2010)
located_in cell surface
inferred from direct assay
(Kirkpatrick et al., 2006)
is_active_in external side of plasma membrane
inferred from direct assay
(Ell et al., 2024)
located_in external side of plasma membrane
inferred from mutant phenotype
(Ayers et al., 2012)
located_in perivitelline space
inferred from high throughput direct assay
(Lye et al., 2014)
Expression Deduced from Reporters
Reporter: P{lacZ}dallyP1
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Nakato et al., 1995)
third instar larval stage
wing disc
(Nakato et al., 1995)
eye-antennal disc
(Nakato et al., 1995)
larval inner optic anlage
(Nakato et al., 1995)
leg disc
(Nakato et al., 1995)
lamina anlage
(Nakato et al., 1995)
Reporter: P{PZ}dally06464
Stage
Tissue/Position (including subcellular localization)
Reference
second instar larval stage -- third instar larval stage
eye-antennal disc
(Zhang et al., 2013)
third instar larval stage
larval inner optic anlage
(Nakato et al., 1995)
lamina anlage
(Nakato et al., 1995)
genital disc | ubiquitous | uniform
(Tsuda et al., 1999)
dorsal-ventral compartment boundary of the wing disc | dorsal to
(Makhijani et al., 2007)
dorsal-ventral compartment boundary of the wing disc | ventral to
(Makhijani et al., 2007)
anterior-posterior compartment boundary of the wing disc | anterior to
(Makhijani et al., 2007)
dorsal-ventral compartment boundary of the haltere disc | dorsal to | restricted

Comment: anterior compartment only

(Makhijani et al., 2007)
dorsal-ventral compartment boundary of the haltere disc | restricted | ventral to

Comment: anterior compartment only

(Makhijani et al., 2007)
adult stage
adult optic lobe
(Vosshall, 1998.8.10)
adult brain | restricted
(Vosshall, 1998.8.10)
adult central complex
(Vosshall, 1998.8.10)
oogenesis
germarium cap cell
(Guo and Wang, 2009)
Reporter: PBac{602.P.SVS-1}dallyCPTI001039
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult optic lobe
(Knowles-Barley, 2011.8.24)
adult gnathal ganglion
(Knowles-Barley, 2011.8.24)
Reporter: PBac{810.P.FSVS-2}dallyCPTI004473
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult optic lobe
(Knowles-Barley, 2011.8.24)
adult brain | restricted
(Knowles-Barley, 2011.8.24)
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\dally 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
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 65 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 28 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of dally
Transgenic constructs containing regulatory region of dally
Aberrations (Deficiencies and Duplications) ( 3 )
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, with Scer\GAL4unspecified
lethal (with dallyf01984)
lethal (with dallySG18)
lethal (with Df(3L)ED4413)
lethal | recessive
lethal - all die before end of larval stage
lethal - all die before end of larval stage | recessive
lethal - all die before end of pupal stage, with Scer\GAL429BD
lethal - all die before end of pupal stage, with Scer\GAL4Act5C.PI
lethal - all die during embryonic stage, with Scer\GAL4Act5C.PI
lethal - all die during embryonic stage, with Scer\GAL4ptc-559.1
partially lethal
partially lethal - majority die
partially lethal - majority die, with Scer\GAL429BD
partially lethal - majority die, with Scer\GAL4Act5C.PI
partially lethal - majority die | recessive
partially lethal - majority live
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4ptc-559.1
viable, with Scer\GAL4tin.CΔ4
Sterility
Allele
female sterile | recessive
fertile
male sterile | recessive
Other Phenotypes
Allele
abnormal cell number | oogenesis, with Scer\GAL4C587
abnormal cell number | oogenesis, with Scer\GAL4tj.PU
abnormal cell number | oogenesis, with Scer\GAL4tsl.B
abnormal cell number | oogenesis (with dally80)
abnormal cell number | oogenesis (with dally06464)
abnormal eclosion rhythm, with Scer\GAL4A9
abnormal immune response
abnormal neuroanatomy
abnormal neuroanatomy, with Scer\GAL4G14, Scer\GAL4how-24B
abnormal neuroanatomy | cell non-autonomous | third instar larval stage, with Scer\GAL4hh-Gal4
abnormal neuroanatomy | embryonic stage, with Scer\GAL4elav.PU
abnormal neuroanatomy | larval stage
abnormal starvation stress response | larval stage
decreased cell death | larval stage, with Scer\GAL4Act.PU
decreased cell death | larval stage, with Scer\GAL4C-765
decreased cell number
decreased size | adult stage
decreased size | larval stage | somatic clone, with Scer\GAL4Act.PU
hyperplasia | adult stage, with Scer\GAL4C587
increased body size, with Scer\GAL429BD
increased cell death | dominant | larval stage
increased cell death | larval stage | somatic clone
increased cell size | embryonic stage
increased occurrence of cell division | adult stage | female, with Scer\GAL4C587
increased occurrence of cell division | spermatogenesis, with Scer\GAL4VP16.nanos.UTR
increased size | larval stage | somatic clone, with Scer\GAL4Act.PU
long lived, with Scer\GAL4elav.PU
segment polarity phenotype
segment polarity phenotype | germline clone
short lived, with Scer\GAL4Ugt36A1.PK
some die during embryonic stage, with Scer\GAL4Act5C.PI
some die during embryonic stage, with Scer\GAL4ptc-559.1
some die during larval stage | recessive
some die during pupal stage, with Scer\GAL429BD
some die during pupal stage, with Scer\GAL4Act5C.PI
some die during third instar larval stage
visible
visible, with Scer\GAL4ci.PU
visible, with Scer\GAL4hh.PU
visible, with Scer\GAL4ptc-559.1
visible, with Scer\GAL4sca-537.4
visible | adult stage
visible | cell non-autonomous, with Scer\GAL4en-GAL4-33
visible | recessive
Phenotype manifest in
Allele
abdominal tergite 6 | female
antenna
anterior crossvein, with Scer\GAL4en-e16E
anterior dorsocentral bristle
anterior dorsocentral bristle (with dally06464)
anterior dorsocentral bristle (with dallygem)
anterior dorsocentral bristle (with dallyΔP-188)
anterior dorsocentral bristle (with dallyΔP-527)
anterior dorsocentral bristle (with Df(3L)Scf-R6)
anterior dorsocentral bristle | absent
axon
central nervous system
chemosensory ventral triple row bristle
costal vein, with Scer\GAL4A9
dendrite
dendritic arborizing neuron
dendritic tree | cell non-autonomous | third instar larval stage, with Scer\GAL4hh-Gal4
denticle
distal dendrite | cell non-autonomous | third instar larval stage, with Scer\GAL4hh-Gal4
dorsal row of triple row of wing sensilla
dorsal row of triple row of wing sensilla (with dally06464)
dorsal row of triple row of wing sensilla (with dallygem)
dorsocentral bristle, with Scer\GAL4ptc-559.1
dorsocentral bristle, with Scer\GAL4sca-537.4
dorsocentral bristle | increased number, with Scer\GAL4A9, dallygem
dorsocentral bristle | increased number, with Scer\GAL4A9, dallyUAS.cJa
dorsocentral bristle | increased number, with Scer\GAL4A9, dallyΔP-527/dallygem
dorsocentral bristle | increased number (with dallygem), with Scer\GAL4A9, dallyUAS.cJa
dorsocentral bristle | increased number (with dallyΔP-527), with Scer\GAL4A9, dallyUAS.cJa
embryo
embryo | germline clone
embryonic/first instar larval cuticle
embryonic epidermis
embryonic epidermis, with Scer\GAL4en-e16E
embryonic epidermis, with Scer\GAL4prd.RG1
embryonic epidermis (with dallygem)
embryonic epidermis (with dallyΔP-188)
embryonic epidermis | germline clone
eye
eye, with Scer\GAL4ey.PH
eye-antennal disc
eye photoreceptor cell, with Scer\GAL4ey.PH
female germline cell | increased number, with Scer\GAL4C587
female germline stem cell
female germline stem cell, with Scer\GAL4bab1-Pgal4-2
female germline stem cell (with dally80)
female germline stem cell (with dally06464)
female germline stem cell | increased number, with Scer\GAL4bab1-Pgal4-2
female germline stem cell | increased number, with Scer\GAL4C587
female germline stem cell | increased number, with Scer\GAL4ptc-559.1
female germline stem cell | increased number, with Scer\GAL4VP16.nanos.UTR
first posterior wing cell, with Scer\GAL4ci.PU
first posterior wing cell, with Scer\GAL4hh.PU
genital disc
genitalia
genitalia (with dallygem)
genitalia (with dallyΔP-188)
germarium, with Scer\GAL4C587
germarium (with dally80)
germarium (with dally06464)
haltere, with Scer\GAL4en-e16E
haltere disc, with Scer\GAL4en-e16E
haltere disc posterior compartment, with Scer\GAL4en-e16E
imaginal disc
lamina
lamina anlage & mitotic cell cycle
larva, with Scer\GAL429BD
larval abdominal segmental nerve, with Scer\GAL4G14, Scer\GAL4how-24B
larval intersegmental nerve branch ISNb of A1-7 | embryonic stage, with Scer\GAL4elav.PU
larval multidendritic class IV neuron | cell non-autonomous | third instar larval stage, with Scer\GAL4hh-Gal4
larval segmental nerve branch SNa of A1-7 | embryonic stage, with Scer\GAL4elav.PU
leg, with Scer\GAL429BD
longitudinal vein
macrochaeta
male genitalia
male genitalia | absent
mechanosensory chaeta
mesothoracic tergum & chaeta | supernumerary, with Scer\GAL4A9
mesothoracic tergum & chaeta | supernumerary, with Scer\GAL4hh-Gal4
microchaeta
NMJ bouton | increased number | larval stage
nucleus | embryonic stage
ommatidium
ommatidium, with Scer\GAL4ey.PH
ovary, with Scer\GAL4C587
ovary | adult stage (with dally80)
ovary | adult stage (with dally06464)
ovary | larval stage, with Scer\GAL4C587
ovary | larval stage, with Scer\GAL4tj.PU
ovary | larval stage, with Scer\GAL4tsl.B
ovary | larval stage (with dally80)
ovary | larval stage (with dally06464)
posterior crossvein, with Scer\GAL471B
posterior crossvein, with Scer\GAL4en-e16E
posterior crossvein, with Scer\GAL4hh-Gal4
posterior dorsocentral bristle
posterior dorsocentral bristle (with dally06464)
posterior dorsocentral bristle (with dallygem)
posterior dorsocentral bristle (with dallyΔP-188)
posterior dorsocentral bristle (with dallyΔP-527)
posterior dorsocentral bristle | increased number, with Scer\GAL429BD
posterior dorsocentral bristle | increased number, with Scer\GAL4A9
posterior dorsocentral bristle | increased number, with Scer\GAL4dpp.blk1
posterior dorsocentral bristle | increased number, with Scer\GAL4en-e16E
posterior dorsocentral bristle | increased number, with Scer\GAL4hh-Gal4
posterior dorsocentral bristle | increased number, with Scer\GAL4ptc-559.1
posterior postalar bristle (with dallygem)
posterior postalar bristle (with dallyΔP-527)
posterior supraalar bristle
posterior supraalar bristle (with dally06464)
posterior supraalar bristle (with dallygem)
posterior supraalar bristle (with dallyΔP-188)
posterior supraalar bristle (with dallyΔP-527)
posterior supraalar bristle (with Df(3L)Scf-R6)
pupa, with Scer\GAL429BD
scutellar bristle | increased number, with Scer\GAL429BD
scutellar bristle | increased number, with Scer\GAL4A9
scutellar bristle | increased number, with Scer\GAL4dpp.blk1
scutellar bristle | increased number, with Scer\GAL4en-e16E
scutellar bristle | increased number, with Scer\GAL4hh-Gal4
scutellar bristle | increased number, with Scer\GAL4ptc-559.1
sensory mother cell (with dallygem)
sensory mother cell (with dallyΔP-527)
sensory mother cell | somatic clone
spermatogonium, with Scer\GAL4VP16.nanos.UTR
taste bristle
wing
wing, with Scer\GAL4A9
wing, with Scer\GAL4dpp.blk1
wing, with Scer\GAL4en-e16E
wing, with Scer\GAL4nub-AC-62
wing, with Scer\GAL4ptc-559.1
wing (with dallygem)
wing (with dallyΔP-527)
wing | distal
wing blade anterior compartment, with Scer\GAL4en-e16E
wing blade anterior compartment | cell non-autonomous, with Scer\GAL4en-GAL4-33
wing blade posterior compartment, with Scer\GAL4en-e16E
wing disc, with Scer\GAL429BD
wing disc, with Scer\GAL4ci.PC
wing disc, with Scer\GAL4en-e16E
wing disc | larval stage
wing disc | larval stage, with Scer\GAL4C-765
wing disc | larval stage | somatic clone
wing disc | larval stage | somatic clone, with Scer\GAL4Act.PU
wing margin
wing margin, with Scer\GAL4hh-Gal4
wing margin bristle
wing margin bristle, with Scer\GAL4nub-AC-62
wing margin bristle | anterior
wing margin bristle | anterior (with dallygem)
wing margin bristle | anterior (with dallyΔP-527)
wing margin bristle | anterior (with Df(3L)Scf-R6)
wing margin bristle | decreased number
wing nerve TSM(1) & sensillum campaniformium (with dallygem)
wing nerve TSM(1) & sensillum campaniformium (with dallyΔP-527)
wing nerve TSM(2) & sensillum campaniformium (with dallygem)
wing nerve TSM(2) & sensillum campaniformium (with dallyΔP-527)
wing sensillum
wing vein
wing vein, with Scer\GAL429BD
wing vein, with Scer\GAL471B
wing vein, with Scer\GAL4A9
wing vein, with Scer\GAL4ptc-559.1
wing vein | ectopic
wing vein | ectopic, with Scer\GAL471B
wing vein | ectopic, with Scer\GAL4A9
wing vein | ectopic, with Scer\GAL4dpp.blk1
wing vein | ectopic, with Scer\GAL4en-e16E
wing vein | ectopic, with Scer\GAL4hh-Gal4
wing vein L4 | ectopic, with Scer\GAL4ci.PC
wing vein L4 | ectopic, with Scer\GAL4en-e16E
wing vein L5
wing vein L5 (with dallygem)
wing vein L5 (with dallyΔP-527)
wing vein L5 (with Df(3L)Scf-R6)
wing vein L5 | somatic clone
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) (7)
Hsap\GPC5
11 of 14
Yes
Yes
4  
Hsap\GPC3
7 of 14
No
Yes
5  
Hsap\GPC1
3 of 14
No
No
Hsap\GPC2
3 of 14
No
No
1  
Hsap\GPC4
3 of 14
No
No
2  
Hsap\GPC6
3 of 14
No
No
1  
Hsap\SYDE1
1 of 14
No
No
1  
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) (7)
Rnor\Gpc3
5 of 14
Yes
Yes
Rnor\Gpc1
3 of 14
No
No
Rnor\Gpc2
3 of 14
No
No
Rnor\Gpc4
3 of 14
No
No
Rnor\Gpc6
3 of 14
No
No
Rnor\Gpc5
2 of 14
No
Yes
Rnor\Syde1
1 of 14
No
No
Mus musculus (laboratory mouse) (7)
Mmus\Gpc5
10 of 14
Yes
Yes
Mmus\Gpc3
7 of 14
No
Yes
Mmus\Gpc1
3 of 14
No
No
Mmus\Gpc2
3 of 14
No
No
Mmus\Gpc4
3 of 14
No
No
Mmus\Gpc6
3 of 14
No
No
Mmus\Syde1
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (5)
Xtro\gpc5
5 of 13
Yes
No
Xtro\gpc3
4 of 13
No
Yes
Xtro\gpc1
2 of 13
No
No
Xtro\gpc4
2 of 13
No
No
Xtro\atp6v1a
1 of 13
No
No
Danio rerio (Zebrafish) (10)
Drer\gpc5a
10 of 14
Yes
Yes
Drer\gpc5b
8 of 14
No
Yes
Drer\gpc3
6 of 14
No
Yes
Drer\gpc5c
6 of 14
No
Yes
Drer\gpc1a
3 of 14
No
No
Drer\gpc1b
3 of 14
No
No
Drer\gpc4
3 of 14
No
No
Drer\gpc6a
3 of 14
No
No
Drer\gpc2
2 of 14
No
No
Drer\gpc6b
2 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (3)
Cele\gpn-1
3 of 14
Yes
No
Cele\lon-2
2 of 14
No
Yes
Cele\C03H12.1
1 of 14
No
Yes
Anopheles gambiae (African malaria mosquito) (2)
Agam\AgaP_AGAP005799
11 of 12
Yes
Yes
Agam\AgaP_AGAP003559
3 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:dally. 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) (1)
dlp
5 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 2 )
    Human Ortholog
    Disease
    Evidence
    References
    GPC3; glypican 3
    model of  Simpson-Golabi-Behmel syndrome type 1
    IEA
    (FlyBase, 2019-)
    model of  nephroblastoma
    IEA
    (FlyBase, 2019-)
    Modifiers Based on Experimental Evidence ( 1 )
    Allele
    Disease
    Interaction
    References
    dallyEP581
    exacerbates  tauopathy
    modeled by Hsap\MAPTV337M.UAS
    (Shulman and Feany, 2003)
    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?
    GPC5; glypican 5
    11 of 14
    602446
         
        GPC3; glypican 3
        7 of 14
        300037
         
        GPC1; glypican 1
        3 of 14
        600395
            GPC2; glypican 2
            3 of 14
            618446
                 
                GPC4; glypican 4
                3 of 14
                300168
                 
                GPC6; glypican 6
                3 of 14
                604404
                 
                Functional Complementation Data
                Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
                Interactions
                Summary of Physical Interactions
                Summary of Genetic Interactions
                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
                dally
                suppressible
                arm
                (Lin and Perrimon, 1999)
                dally
                suppressible
                CycA
                (Nakato et al., 2002)
                dally
                suppressible
                Dad
                (Guo and Wang, 2009)
                dally
                suppressible
                dlp
                (Nakato et al., 2024)
                dally
                enhanceable
                dlp
                (Nakato et al., 2024, Baron et al., 2009)
                dally
                suppressible
                dpp
                (Liu et al., 2024, Jackson et al., 1997)
                dally
                enhanceable
                dpp
                (Jackson et al., 1997)
                dally
                suppressible
                fwe
                (Merino et al., 2022)
                dally
                suppressible
                Kr
                (Liu et al., 2024)
                dally
                suppressible
                Mad
                (Kirkpatrick et al., 2006)
                dally
                suppressible
                nord
                (Akiyama et al., 2022)
                dally
                enhanceable
                nord
                (Akiyama et al., 2022)
                dally
                suppressible
                ptc
                (Takeo et al., 2005)
                dally
                enhanceable
                put
                (Fujise et al., 2001)
                dally
                enhanceable
                Sdc
                (Poe et al., 2017)
                dally
                enhanceable
                sfl
                (Lin and Perrimon, 1999)
                dally
                enhanceable
                sll
                (Luders et al., 2003)
                dally
                suppressible
                tkv
                (Guo and Wang, 2009)
                dally
                enhanceable
                wdp
                (Koh et al., 2023)
                dally
                suppressible
                wg
                (Tsuda et al., 1999)
                dally
                enhanceable
                wg
                (Fujise et al., 2001, Lin and Perrimon, 1999, Tsuda et al., 1999)
                Starting gene(s)
                Interaction type
                Interacting gene(s)
                Reference
                dlp
                enhanceable
                dally
                (Franch-Marro et al., 2005)
                dpp
                suppressible
                dally
                (Jackson et al., 1997)
                fz2
                enhanceable
                dally
                (Lin and Perrimon, 1999)
                fz2
                suppressible
                dally
                (Lin and Perrimon, 1999)
                hh
                suppressible
                dally
                (Jackson et al., 1997)
                L
                enhanceable
                dally
                (Singh et al., 2005)
                Notum
                suppressible
                dally
                (Kakugawa et al., 2015)
                Sdc
                enhanceable
                dally
                (Poe et al., 2017)
                Sema1a
                enhanceable
                dally
                (Cho et al., 2012)
                sfl
                suppressible
                dally
                (Weigelt et al., 2020)
                stet
                suppressible
                dally
                (Liu et al., 2010)
                Sulf1
                suppressible
                dally
                (You et al., 2011, Kleinschmit et al., 2010)
                wg
                suppressible
                dally
                (Moline et al., 2000, Tsuda et al., 1999)
                wg
                enhanceable
                dally|fz2
                (Moline et al., 2000)
                wg
                enhanceable
                dally
                (Moline et al., 2000, Tsuda et al., 1999)
                External Data
                Subunit Structure (UniProtKB)
                Interacts with nord; the interaction promotes dally degradation (PubMed:35609633). Interacts with Magu (PubMed:35301437). As part of the dally/ Magu complex, associates with fwe (isoforms ubi, LoseA and LoseB) and is unable to interact with fwe independently of Magu (PubMed:35301437).
                (UniProt, Q24114 )
                Linkouts
                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
                Class of Gene
                Genomic Location and Detailed Mapping Data
                Chromosome (arm)
                3L
                Recombination map
                3-27
                Cytogenetic map
                66E1-66E3
                Sequence location
                FlyBase Computed Cytological Location
                Cytogenetic map
                Evidence for location
                66E1-66E3
                Limits computationally determined from genome sequence between P{PZ}l(3)0162901629&P{PZ}mRpL1210534 and P{EP}Hsp26EP3336&P{EP}Hsp26EP3315
                Experimentally Determined Cytological Location
                Cytogenetic map
                Notes
                References
                66E-66E
                (determined by in situ hybridisation)
                (Merabet et al., 2002)
                66E1-66E2
                (determined by in situ hybridisation)
                (Abdelilah-Seyfried et al., 2000, Spradling et al., 1999)
                66D-66E
                (Nakato, 1995.7.20)
                66E1-66E2
                (BDGP Project Members, 1994-1999)
                Experimentally Determined Recombination Data
                Location

                3-27

                (FlyBase, 2016)

                3-21.1

                (Comeron et al., 2012)

                3-

                (Nakato, 1995.7.20, Chorsky and Belote, 1994)

                3-31.9

                Left of (cM)
                Right of (cM)
                Notes

                Mapped by meiotic recombination to a position very near h.

                (Chorsky and Belote, 1994)
                Stocks and Reagents
                Stocks (126)
                Genomic Clones (63)
                cDNA Clones (219)
                 

                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
                Antibody Information
                Laboratory Generated Antibodies
                 

                polyclonal

                (Ayers et al., 2012, Tsuda et al., 1999)
                Commercially Available Antibodies
                 
                Cell Line Information
                Publicly Available Cell Lines
                 
                  Other Stable Cell Lines
                   
                    Other Comments

                    Studies with a dally transgene that cannot be modified with heparan sulfate shows that the function of the dally product does not depend entirely on its heparan sulfate modification.

                    (Kirkpatrick et al., 2006)

                    dally in combination with dlp is required for hh protein movement during wing disc development.

                    (Han et al., 2004)

                    One of 42 Drosophila genes identified as being most likely to reveal molecular and cellular mechanisms of nervous system development or plasticity relevant to human Mental Retardation disorders.

                    (Inlow and Restifo, 2004)

                    dally and dlp are dispensable for wg signalling in embryos.

                    (Desbordes and Sanson, 2003)

                    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)

                    dally mutant cell division phenotypes are selectively rescued by reduction in CycA function. dally mutants show abnormal CycA protein levels in the dividing cells affected, with appreciable levels of CycA protein remaining in late prophase and metaphase, stages where CycA protein is normally absent.

                    (Nakato et al., 2002)

                    dally appears to be an important regulator of sensory organ formation via wg and dpp mediated specification of proneural clusters.

                    (Fujise et al., 2001)

                    dally expression is controlled by a TATA-less promoter. dally expression appears to be regulated posttranscriptionally by modulating translation efficiency and mRNA stability.

                    (Tsuda et al., 2001)

                    dally may act as a co-receptor for wg, and together with fz2 modulates both short- and long-range activities of wg.

                    (Lin and Perrimon, 1999)

                    Candidate gene for quantitative trait locus.

                    (Macdonald and Goldstein, 1999)

                    dally can differentially influence signalling mediated by two growth factors, and may form a regulatory component of both wg and dpp receptor complexes.

                    (Tsuda et al., 1999)

                    Genetic experiments suggest that dally is haplolethal.

                    (Tsuda et al., 1999)

                    dally is required for normal dpp signaling during development.

                    (Jackson et al., 1997)

                    dally mutants show decreased viability and morphological defects in several adult tissues including the eye, antenna, wing and genitalia. The lethality and dally phenotypes are partially suppressed by deleting one copy of CycA.

                    (Futch et al., 1995)

                    dally is required for ordered cell division in the larval brain and eye disc, mutations affect the timing of the G2-M transition.

                    (Futch et al., 1995)

                    Mutants show specific cell division defects in these tissues in the absence of any gross morphological disorganisation.

                    (Nakato et al., 1995)

                    Molecular cloning of dally cDNA shows it encodes a putative integral membrane proteoglycan of the Glypican family.

                    (Nakato et al., 1995)

                    dally is required for the normal patterning of cell division in at least two larval tissues, the eye disc and lamina.

                    (Nakato et al., 1995)

                    dally was initially identified as a P-element enhancer trap insertion with expression in G2 and M phases of the first LPC division, expression overlaps with that of CycB. dally mutants also show abnormalities in the eye, antenna, wing and genitalia.

                    (Selleck et al., 1994)

                    dally is required for the ordered progression of both lamina precursor cell (LPC) divisions before they differentiate into neurons. An analysis of LPC cell cycle progression suggests primary failure is a delay in the entry into the first M phase from G2.

                    (Selleck et al., 1994)
                    Relationship to Other Genes
                    Source for database merge of

                    Source for merge of: dally gem

                    (Selleck, 1999.11.2)

                    Source for merge of: dally l(3)SG18

                    (Haerry, 2011.12.5)
                    Additional comments
                    Nomenclature History
                    Source for database identify of
                    Nomenclature comments
                    Etymology

                    Gene named on the basis of the cell division defects seen in mutants.

                    (Nakato et al., 1995)
                    Synonyms and Secondary IDs (13)
                    Reported As
                    Symbol Synonym
                    CG4974
                    (Yamamoto-Hino et al., 2015, Mosallanejad et al., 2010, Ni et al., 2010.12.1, Perkins et al., 2009, Norga et al., 2003, Mount, 2001.8.14, Staatz, 2000.12.21)
                    CT15952
                    (Hynes and Zhao, 2000)
                    Dally
                    (St Johnston, 2023, Gradilla and Guerrero, 2022, Montanari et al., 2022, Niu and Liu, 2022, Madamanchi et al., 2021, Vidaurre and Chen, 2021, Gou et al., 2020, Khan et al., 2020, Rust et al., 2020, Zhang and Cai, 2020, Fu et al., 2017, Koch et al., 2017, Lee et al., 2016, Dorn and Dorn, 2015, Yamamoto-Hino et al., 2015, Bausek, 2013, Gradilla and Guerrero, 2013, Mbodj et al., 2013, Muha and Müller, 2013, Umulis and Othmer, 2013, Matunis et al., 2012, Raftery and Umulis, 2012, Ben-Zvi et al., 2011, Eliazer and Buszczak, 2011, Hamaratoglu et al., 2011, Harris and Ashe, 2011, You et al., 2011, Serpe et al., 2008, Sexton and Harrison, 2008, Vyas et al., 2008, Wang et al., 2008, Hufnagel et al., 2007, Clevers, 2006, Wilson and Chuang, 2006, Eugster and Eaton, 2005, Kirkbride, 2005, Peifer, 1999, Bassett et al., 1997)
                    anon-EST:Liang-1.63
                    clone 1.63
                    (Liang and Biggin, 1998)
                    dally
                    (Li et al., 2025, Mendoza Andrade et al., 2025, Cui et al., 2024, Ell et al., 2024, Eslahi et al., 2024, Jiménez-Jiménez et al., 2024, Li et al., 2024, Pignataro et al., 2024, Simon et al., 2024, Ascencio et al., 2023, Golubev et al., 2023, Gude et al., 2023, Aguirre-Tamaral et al., 2022, Akiyama et al., 2022, Li et al., 2022, Liu et al., 2022, Marcogliese et al., 2022, Merino et al., 2022, Perlegos et al., 2022, Rahman et al., 2022, Romanova-Michaelides et al., 2022, Waghmare and Page-McCaw, 2022, Guzman et al., 2021, Hagen et al., 2021, Park et al., 2021, Ramanujam et al., 2021, Hayashi et al., 2020, McGough et al., 2020, Rushton et al., 2020, Statzer and Ewald, 2020, Wei et al., 2020, Weigelt et al., 2020, Zhao et al., 2020, Brann et al., 2019, Cai et al., 2019, Drummond-Barbosa, 2019, Hu et al., 2019, Meltzer et al., 2019, Nelson et al., 2019, Croset et al., 2018, Kanai et al., 2018, Kang et al., 2018, Lee et al., 2018, Malzer et al., 2018, Mehrotra and Deshpande, 2018, Tseng et al., 2018, Aw et al., 2017, Barrio and Milán, 2017, Fochler et al., 2017, González-Méndez et al., 2017, Poe et al., 2017, Revaitis et al., 2017, Rohde et al., 2017, Upadhyay et al., 2017, Barton et al., 2016, Clandinin and Owens, 2016-, Gene Disruption Project members, 2016-, Levings et al., 2016, Nakato and Li, 2016, Norman et al., 2016, Yamazaki et al., 2016, Dembeck et al., 2015, Duff et al., 2015, Grotewiel and Bettinger, 2015, Kakugawa et al., 2015, Kim et al., 2015, Lu et al., 2015, Simon and Guerrero, 2015, Gersten et al., 2014, Ikmi et al., 2014, Merlo et al., 2014, Avanesov and Blair, 2013, Bilioni et al., 2013, Curtis et al., 2013, Dejima et al., 2013, Khan et al., 2013, Kleinschmit et al., 2013, Sen et al., 2013, Webber et al., 2013, Xin et al., 2013, Zhang et al., 2013, Zhu and Zhang, 2013, Avanesov et al., 2012, Ayers et al., 2012, Chen et al., 2012, Cho et al., 2012, Hayashi et al., 2012, Hazelett et al., 2012, Sánchez-Hernández et al., 2012, Xia et al., 2012, Carreira et al., 2011, Chang et al., 2011, Dejima et al., 2011, Eliazer and Buszczak, 2011, Eliazer et al., 2011, Haerry, 2011.12.5, Kirilly et al., 2011, Losick et al., 2011, Slattery et al., 2011, Toku et al., 2011, Turner et al., 2011, Wang et al., 2011, Wojcinski et al., 2011, You et al., 2011, Zhang et al., 2011, Ayers et al., 2010, Kleinschmit et al., 2010, Knowles-Barley et al., 2010, Liu et al., 2010, Vuilleumier et al., 2010, Williams et al., 2010, Wu et al., 2010, Yan et al., 2010, Baron et al., 2009, Guo and Wang, 2009, Hayashi et al., 2009, Hiller et al., 2009, Akiyama et al., 2008, Baena-Lopez et al., 2008, Bornemann et al., 2008, Buszczak et al., 2007, Crickmore and Mann, 2007, Crickmore and Mann, 2007, Eugster et al., 2007, Grueber et al., 2007, Kaplow et al., 2007, Makhijani et al., 2007, Yan and Lin, 2007, Zeitouni et al., 2007, de Navas et al., 2006, Kirkpatrick et al., 2006, Wendler et al., 2006, Yao et al., 2006, Zhang et al., 2006, Briscoe and Therond, 2005, Eldar and Barkai, 2005, Glise et al., 2005, Häcker et al., 2005, Liu et al., 2005, Strigini, 2005, Torroja et al., 2005, Kreuger et al., 2004, Tsuda et al., 1999)
                    gem
                    (Macdonald and Goldstein, 1999, Chorsky and Belote, 1994)
                    l(3)06464
                    l(3)SG18
                    (Haerry, 2011.12.5)
                    Name Synonyms
                    Dally
                    (Daly et al., 2022, Yang et al., 2021, Mii and Takada, 2020, Nishihara, 2020, Nagarajan et al., 2015, Shimmi and Newfeld, 2013, Niehrs, 2012, Ben-Zvi et al., 2011, Choo et al., 2011, Akiyama et al., 2008, Williams et al., 2008, Kerszberg and Wolpert, 2007)
                    Division abnormally delayed
                    (Eliazer and Buszczak, 2011, Harris and Ashe, 2011, Bornemann et al., 2008)
                    dally
                    (Titlow et al., 2023, Jonchère et al., 2017, Levings et al., 2016, Neto et al., 2016, Avanesov and Blair, 2013, Guo and Wang, 2009, Yamashita et al., 2004, Tsuda et al., 1999)
                    division abnormally delayed
                    (Upadhyay et al., 2017, Zhang et al., 2013, Eliazer and Buszczak, 2011, You et al., 2011, Wu et al., 2010, Kaplow et al., 2007, de Navas et al., 2006, Burnette et al., 2005, Nakato et al., 1995)
                    genitalia missing
                    (Chorsky and Belote, 1994)
                    lethal (3) SG18
                    Secondary FlyBase IDs
                    • FBgn0011577
                    • FBgn0010889
                    • FBgn0014962
                    • FBgn0002464
                    Datasets (0)
                    Study focus (0)
                    Experimental Role
                    Project
                    Project Type
                    Title
                    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
                    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
                    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
                    DroID - A comprehensive database of gene and protein interactions.
                    FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
                    FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                    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.
                    Reactome Metabolic Pathways - An open-source, open access, manually curated and peer-reviewed metabolic pathway database.
                    SignaLink - A signaling pathway resource with multi-layered regulatory networks.
                    References (391)