FB2026_02 , released June 18, 2026
Gene: Dmel\upd1
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General Information
Symbol
Dmel\upd1
Species
D. melanogaster
Name
unpaired 1
Annotation Symbol
CG5993
Feature Type
protein_coding_gene
FlyBase ID
FBgn0004956
Gene Model Status
Current
Stock Availability
33 publicly available
Gene Summary
unpaired 1 (upd1) encodes a secreted glycoprotein that is able to act at a distance as the primary ligand for the JAK/STAT signaling pathway. Although semi-redundant with upd2 and upd3, upd1 depletion results in embryonic lethality with an atypical gap gene-like segmentation phenotype. [Date last reviewed: 2019-03-21] (FlyBase Gene Snapshot)
All Summaries
Also Known As

upd, unpaired, os, sisC, outstretched

Key Links
Genomic Location
Cytogenetic map
17A5-17A5
Sequence location
Recombination map
1-60
RefSeq locus
NC_004354 REGION:18305354..18309291
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 (34 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
enables cytokine activity
inferred from direct assay
(Wright et al., 2011)
inferred from genetic interaction with FLYBASE:dome; FB:FBgn0043903
(Hombria et al., 2005)
enables extracellular matrix binding
inferred from direct assay
(Hombria et al., 2005)
enables receptor ligand activity
inferred from direct assay
(Devergne et al., 2007)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (29 terms)
Terms Based on Experimental Evidence (26 terms)
CV Term
Evidence
References
involved_in apical constriction
inferred from mutant phenotype
(Bertet et al., 2009)
involved_in border follicle cell migration
inferred from mutant phenotype
(Xi et al., 2003)
involved_in cell surface receptor signaling pathway via JAK-STAT
inferred from direct assay
(Shen et al., 2022, Wright et al., 2011, Bach et al., 2007, Devergne et al., 2007, Hombria et al., 2005, Muller et al., 2005, Harrison et al., 1998)
inferred from mutant phenotype
(Osman et al., 2012, Wright et al., 2011, Wawersik et al., 2005, Johansen et al., 2003, Chen et al., 2002, Sefton et al., 2000, Zeidler et al., 1999)
inferred from genetic interaction with FLYBASE:hop; FB:FBgn0004864
(Wright et al., 2011)
inferred from genetic interaction with FLYBASE:Stat92E; FB:FBgn0016917
(Wright et al., 2011)
inferred from genetic interaction with FLYBASE:Socs36E; FB:FBgn0041184
(Wright et al., 2011)
inferred from genetic interaction with FLYBASE:dome; FB:FBgn0043903
(Wright et al., 2011)
inferred from genetic interaction with FLYBASE:Ptp61F; FB:FBgn0267487
(Wright et al., 2011)
involved_in compound eye morphogenesis
inferred from mutant phenotype
(Tsai and Sun, 2004)
involved_in defense response to Gram-negative bacterium
inferred from high throughput mutant phenotype
(Cronin et al., 2009)
involved_in determination of digestive tract left/right asymmetry
inferred from mutant phenotype
(Lai et al., 2023)
involved_in establishment of ommatidial planar polarity
inferred from mutant phenotype
(Zeidler et al., 1999)
involved_in eye-antennal disc morphogenesis
inferred from mutant phenotype
(Ekas et al., 2006)
involved_in follicle cell of egg chamber development
inferred from mutant phenotype
(Medioni and Noselli, 2005, Xi et al., 2003)
involved_in foregut morphogenesis
inferred from mutant phenotype
(Josten et al., 2004)
involved_in germ-line stem-cell niche homeostasis
inferred from genetic interaction with FLYBASE:Imp; FB:FBgn0285926
(Toledano et al., 2012)
involved_in hindgut morphogenesis
inferred from mutant phenotype
(Johansen et al., 2003)
involved_in intestinal epithelial structure maintenance
inferred from mutant phenotype
(Osman et al., 2012)
involved_in intestinal stem cell homeostasis
inferred from mutant phenotype
(Guo et al., 2013, Osman et al., 2012)
involved_in locomotor rhythm
inferred from mutant phenotype
(Luo and Sehgal, 2012)
involved_in long-term memory
inferred from mutant phenotype
(Copf et al., 2011)
involved_in male germ-line stem cell population maintenance
inferred from mutant phenotype
(Parrott et al., 2011)
inferred from genetic interaction with UniProtKB:A1YK24,UniProtKB:Q9VCH5,UniProtKB:Q9VCH5
(Parrott et al., 2011)
involved_in mediolateral intercalation
inferred from mutant phenotype
(Bertet et al., 2009)
involved_in negative regulation of innate immune response
inferred from high throughput mutant phenotype
(Cronin et al., 2009)
involved_in paracrine signaling
inferred from direct assay
(Wright et al., 2011)
involved_in primary sex determination
inferred from mutant phenotype
(Jinks et al., 2000, Sefton et al., 2000)
involved_in regulation of embryonic cell shape
inferred from mutant phenotype
(Bertet et al., 2009)
involved_in stem cell division
inferred from mutant phenotype
(Tulina and Matunis, 2001)
involved_in synaptic target recognition
inferred from mutant phenotype
(Kurusu et al., 2008)
involved_in tissue regeneration
inferred from mutant phenotype
(Harris et al., 2020)
involved_in wing disc morphogenesis
inferred from mutant phenotype
(Recasens-Alvarez et al., 2017)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
involved_in blastoderm segmentation
traceable author statement
(Luo and Dearolf, 2001)
involved_in cell surface receptor signaling pathway via JAK-STAT
inferred from electronic annotation with InterPro:IPR031901
(InterPro Project Members, 2004-)
involved_in embryonic development via the syncytial blastoderm
inferred from electronic annotation with InterPro:IPR031901
(InterPro Project Members, 2004-)
involved_in sex determination, establishment of X:A ratio
non-traceable author statement
(Schutt and Nothiger, 2000)
Cellular Component (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in apical part of cell
inferred from direct assay
(Van De Bor et al., 2011)
located_in extracellular region
inferred from direct assay
(Van De Bor et al., 2011, Wright et al., 2011, Hombria et al., 2005, Harrison et al., 1998)
Terms Based on Predictions or Assertions (0 terms)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
unpaired 1 (upd1) encodes a secreted glycoprotein that is able to act at a distance as the primary ligand for the JAK/STAT signaling pathway. Although semi-redundant with upd2 and upd3, upd1 depletion results in embryonic lethality with an atypical gap gene-like segmentation phenotype. [Date last reviewed: 2019-03-21]
Gene Group (FlyBase)
UNPAIRED FAMILY -
The Unpaired (UPD) family encodes the only known cytokines capable of activating the JAK/STAT pathway in D.mel, by binding the receptor dome. This pathway is involved in cell proliferation, embryonic development and stem cell maintenance. (Adapted from FBrf0232479, FBrf0213113 and FBrf0219535).
Pathway (FlyBase)
JAK-STAT SIGNALING PATHWAY CORE COMPONENTS -
The JAK-STAT signaling pathway is initiated by the binding of an extracellular ligand to a cell surface receptor leading to receptor dimerization and the intracellular activation of a Janus kinase (JAK) family member. JAK phosphorylates cytoplasmic STAT family members which dimerize, translocate into the nucleus and regulate target gene expression. In Drosophila, the core pathway is limited to three ligands (the Unpaired family of cytokines), a single receptor (dome), JAK kinase (hop) and STAT (Stat92E). (Adapted from FBrf0225259).
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
os: outstretched small eye
Wings held virtually at right angles to body. Eyes small and rounded. os/oso has wing effect but eyes normal. os/oss has eye effect but wings normal. RK1.
osbdw: outstretched small eye-bending wings
Wings divergent and drooping, size and shape normal. Males sterile. RK2A.
oso: outstretched small eye-outstretched
thumb
oso: outstretched small eye-outstretched
Edith M. Wallace, unpublished.
Wings extremely divergent, often at right angles to body. oso/oss is wild type. RK1.
oss: outstretched small eye-small eye
Eyes small and rounded, high on the head but not bulging. RK1.
upd: unpaired
Embryonic lethal. Mutants show variable larval cuticular phenotype with defects predominantly in the mesothorax and the fifth abdominal segment (Nusslein-Volhard et al., 1984; Gergen and Wieschaus, 1986); also some head defects and fourth, sixth, seventh, and eighth abdominal segment defects. Heterozygotes show 88-93% viability (Wieschaus and Noell, 1986).
Summary (Interactive Fly)

interleukin-like ligands of Domeless - activators of JAK/STAT signaling pathway - mutation results in the stripe-specific loss of expression of even-skipped, fushi tarazu, and runt

Gene Model and Products
Number of Transcripts
3
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\upd1 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 Q9VWW4)

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 5.52

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

Gene model reviewed during 5.43

Gene model reviewed during 5.42

Gene model reviewed during 5.56

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
upd1-RA
FBtr0074581
NM_080356
2642
413
upd1-RB
FBtr0112847
NM_001103545
2443
413
upd1-RC
FBtr0343592
NM_001298473
2602
413
Additional Transcript Data and Comments
Reported size (kB)

2.2 (northern blot)

(Harrison et al., 1998)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
upd1-PA
FBpp0074353
47.0
413
12.00
Q9VWW4
NP_525095
AAF48823
upd1-PB
FBpp0111760
47.0
413
12.00
Q9VWW4
NP_001097015
ABW09444
upd1-PC
FBpp0310189
47.0
413
12.00
Q9VWW4
NP_001285402
AHN59872
Polypeptides with Identical Sequences

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

413 aa isoforms: upd1-PA, upd1-PB, upd1-PC
Additional Polypeptide Data and Comments
Reported size (kDa)

413 (aa); 47 (kD predicted)

(Harrison et al., 1998)
Comments
External Data
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\upd1 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).

4.20

Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 4 -- 5
organism | pair rule expression pattern
(Harrison et al., 1998)
embryonic stage 6 -- 7
organism | segment polarity expression pattern
(Harrison et al., 1998)
embryonic stage 9 -- 17
tracheal pit
(Harrison et al., 1998)
distribution deduced from reporter (Gal4 UAS)
Stage
Tissue/Position (including subcellular localization)
Reference
wandering third instar larval stage
wing hinge primordium | restricted
(Wang et al., 2014)
dorsal mesothoracic pleura primordium | restricted
(Wang et al., 2014)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 4 -- 5
organism | 20-80% egg length
(Savant-Bhonsale et al., 1999)
organism | anterior | dorsal
(Savant-Bhonsale et al., 1999)
embryonic stage 9
ventral ectoderm | striped
(Johnson et al., 2011)
embryonic stage 10 -- 11
organism | segmentally repeated | striped
(Savant-Bhonsale et al., 1999)
embryonic stage 10 -- 12
embryonic hindgut
(Johnson et al., 2011)
presumptive embryonic/larval tracheal system
(Johnson et al., 2011)
embryonic stage 11
small intestine primordium
(Uddin et al., 2011)
embryonic stage 12 -- 14
embryonic small intestine
(Uddin et al., 2011)
embryonic stage 13
small intestine primordium
(Iwaki et al., 2001)
embryonic stage 13 -- 14
imaginal tissue
(Savant-Bhonsale et al., 1999)
embryonic stage 14
ectoderm of pharynx
(Hombria et al., 2005)
ectoderm of hindgut
(Hombria et al., 2005)
embryonic stage 16
gonad
(Brown et al., 2006)
embryonic stage 17
hub cell
(Sheng et al., 2009)
gonad | anterior
(Sheng et al., 2009)
second instar larval stage
wing disc | distal
(Recasens-Alvarez et al., 2017)
late third instar larval stage
lamina furrow
(Wang et al., 2011)
adult stage
adult mushroom body
(Copf et al., 2011)
Additional Descriptive Data

upd1 transcript is expressed in the proximal fold, and two domains of the medial fold, of the wing hinge primordium, and in a small domain corresponding to the ventral body wall.

(Wang et al., 2014)

Brief segmental upd1 expression is detected in the ventral ectoderm at embryonic stage 9. In stages 10-12, upd1 is expressed in the developing tracheal system and in the hindgut and is not longer seen in the ventral ectoderm. No expression is observed in the dorsal ectoderm or dorsal mesoderm.

(Johnson et al., 2011)

upd1 localizes to hub-proximal germ cells in the anterior half of the gonad in stage 17 embryos.

(Sheng et al., 2009)

os transcripts become abundant shortly before cellularization at which time they are present in the trunk region and in an incomplete head stripe but not in the termini. At cellularization, the trunk region expression resolves into 7 stripes. In early gastrulation, 14 stripes appear. In later stages, expression is largely restricted to the tracheal pits.

(Harrison et al., 1998)
Marker for
 
embryonic small intestine
(Uddin et al., 2011)
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage | female
germarium cap cell
(Hayashi et al., 2012)
terminal filament
(Hayashi et al., 2012)
adult stage
calyx of adult mushroom body
(Copf et al., 2011)
adult brain
(Copf et al., 2011)
oogenesis stage S2 -- S6
stalk follicle cell
(Hayashi et al., 2012)
polar follicle cell
(Hayashi et al., 2012)
oogenesis stage S4 -- S10
oocyte associated follicular epithelium | gradient
(Hayashi et al., 2012)
Additional Descriptive Data

os protein localizes to the extracellular matrix.

(Hayashi et al., 2012, Harrison et al., 1998)

upd1 protein is present in the mushroom bodies and accumulates in the calyx, although it covers a broader brain region.

(Copf et al., 2011)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in apical part of cell
inferred from direct assay
(Van De Bor et al., 2011)
located_in extracellular region
inferred from direct assay
(Van De Bor et al., 2011, Wright et al., 2011, Hombria et al., 2005, Harrison et al., 1998)
Expression Deduced from Reporters
Reporter: P{GawB}E132
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
leg disc | restricted
(Halder et al., 1995)
wing disc | restricted
(Halder et al., 1995)
eye-antennal disc | restricted
(Halder et al., 1995)
imaginal disc | restricted
(Halder et al., 1995)
haltere disc | restricted
(Halder et al., 1995)
second instar larval stage
eye-antennal disc | ventral
(Pignoni and Zipursky, 1997)
wing disc | distal
(Recasens-Alvarez et al., 2017)
second instar larval stage -- early third instar larval stage
larval outer optic anlage | restricted
(Wang et al., 2011)
third instar larval stage
embryonic/larval optic stalk | vicinity of
(Pignoni and Zipursky, 1997)
eye disc | posterior
(Pignoni and Zipursky, 1997)
antennal disc
(Pignoni and Zipursky, 1997)
late third instar larval stage
lamina furrow
(Wang et al., 2011)
wandering third instar larval stage
wing hinge primordium
(Bardot et al., 2002)
adult stage
adult brain | restricted
(Luo and Sehgal, 2012)
adult DN period neuron
(Luo and Sehgal, 2012)
enteroblast
(Osman et al., 2012)
intestinal stem cell
(Osman et al., 2012)
oogenesis
polar follicle cell
(Assa-Kunik et al., 2007)
spermatogenesis
hub cell
(Voog et al., 2008)
adult stage
alpha/beta Kenyon cell | subset
(Copf et al., 2011)
gamma Kenyon cell | subset
(Copf et al., 2011)
mushroom body pedunculus
(Copf et al., 2011)
pharate adult stage P8 -- P9
wing pouch
(Hatini et al., 2013)
mesonotum | lateral
(Hatini et al., 2013)
pharate adult stage P8 -- P15
wing hinge
(Hatini et al., 2013)
mesonotum | restricted
(Hatini et al., 2013)
adult stage
calyx of adult mushroom body
(Copf et al., 2011)
Reporter: P{lacW}PD
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Sun et al., 1995)
third instar larval stage
eye disc | restricted
(Sun et al., 1995)
leg disc | restricted
(Sun et al., 1995)
adult stage
intestinal stem cell of posterior adult midgut epithelium
(Liu et al., 2010)
enteroblast of posterior adult midgut epithelium
(Liu et al., 2010)
oogenesis stage S9 & adult stage | female
polar follicle cell
(Lucas et al., 2013)
Reporter: P{upd1-GAL4.U}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 17
gonadal sheath proper primordium
(Le Bras and Van Doren, 2006)
adult stage
hub cell
(Le Bras and Van Doren, 2006)
adult stage | male
hub cell
(Lee et al., 2016)
adult stage | male & spermatogenesis
hub cell
(Fairchild et al., 2016)
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\upd1 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 ( 19 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 22 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of upd1
Transgenic constructs containing regulatory region of upd1
Aberrations (Deficiencies and Duplications) ( 25 )
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(1)os1A2)
lethal | recessive
lethal - all die before end of larval stage, with Spyo\Cas9dCas9::VPR.wg-ΔC1
lethal - all die before end of larval stage, with Spyo\Cas9dCas9::VPR.αTub84B
lethal - all die before end of P-stage, with Spyo\Cas9dCas9::VPR.foxo
lethal - all die before end of pupal stage
lethal - all die during embryonic stage
lethal - all die during embryonic stage | recessive
partially lethal
partially lethal - majority die | female, with Spyo\Cas9dxCas9-3.7::VPR.αTub84B
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable | male, with Spyo\Cas9dxCas9-3.7::VPR.αTub84B
Sterility
Allele
female sterile
fertile
male sterile
Other Phenotypes
Allele
abnormal axis specification | embryonic stage
abnormal cell number | oogenesis
abnormal cell number | spermatogenesis, with Scer\GAL4tj.PU
abnormal cell shape | adult stage, with Scer\GAL4esg.PU
abnormal cell shape | extended germ band stage
abnormal cell shape | oogenesis
abnormal developmental rate
abnormal developmental rate, with Scer\GAL4P0206
abnormal feeding behavior | adult stage, with Scer\GAL4elav.PU
abnormal immune response | adult stage, with Scer\GAL4NP5130
abnormal immune response | adult stage, with Scer\GAL4r4
abnormal immune response | conditional, with Scer\GAL4NP0001
abnormal locomotor rhythm, with Scer\GAL4mir-279.PC
abnormal locomotor rhythm, with Scer\GAL4tim.PE
abnormal locomotor rhythm, with Scer\GAL4tim.UAS
abnormal locomotor rhythm | RU486 conditional, with Scer\GAL4elav.Switch.PO
abnormal memory | adult stage | RU486 conditional, with Scer\GAL4Mef2.247.Switch
abnormal planar polarity | cell non-autonomous | somatic clone, with Scer\GAL4Act5C.PI
abnormal planar polarity | recessive | somatic clone
abnormal planar polarity | somatic clone, with Scer\GAL430A
abnormal size, with Scer\GAL4en-e16E
abnormal size, with Scer\GAL4ey.PH
abnormal size, with Scer\GAL4GMR.PU
abnormal size | adult stage
abnormal size | adult stage, with Df(1)os1A2
abnormal size | adult stage, with Scer\GAL4bi-omb-Gal4
abnormal size | adult stage, with Scer\GAL4dpp.blk1
abnormal size | adult stage, with Scer\GAL4ey.PH
abnormal size | adult stage, with Scer\GAL4GMR.PF
abnormal size | adult stage, with Scer\GAL4VP16.nanos.UTR
abnormal size | adult stage, with upd1GMR.PB
abnormal size | adult stage, with upd1YC43
abnormal size | adult stage, with upd1YM55
abnormal size | third instar larval stage, with Scer\GAL4dpp.blk1
abnormal size | third instar larval stage, with Scer\GAL4ey.PH
abnormal size | third instar larval stage, with Scer\GAL4ptc.PU
decreased cell number | adult stage | somatic clone, with Scer\GAL4NP5130
decreased occurrence of cell division | adult stage, with Scer\GAL4esg.PU
decreased occurrence of cell division | adult stage, with Scer\GAL4NP5130
decreased occurrence of cell division | third instar larval stage, with Scer\GAL4GMR.PF
decreased size | adult stage | somatic clone, with Scer\GAL4R42A07.-FRT
hyperplasia, with Scer\GAL4C587
hyperplasia, with Scer\GAL4ptc-559.1
hyperplasia | adult stage, with Scer\GAL4dpp.blk1
hyperplasia | adult stage, with Scer\GAL4GMR.PF
hyperplasia | adult stage | somatic clone, with Scer\GAL4Act5C.PI
increased cell growth rate | somatic clone, with Scer\GAL4FRT.Rnor\Cd2.Act5C
increased cell number
increased cell number, with Scer\GAL4GMR.PF
increased cell number | adult stage, with Scer\GAL4esg.PU
increased cell number | adult stage, with Scer\GAL4GMR.PF
increased cell number | adult stage, with Scer\GAL4NP5130
increased cell number | adult stage | male, with Scer\GAL4VP16.nanos.UTR
increased cell number | larval stage, with Scer\GAL4Hand.Δ
increased cell number | larval stage, with Scer\GAL4NP1029
increased cell number | oogenesis, with Scer\GAL4c306
increased cell number | oogenesis, with Scer\GAL4VP16.nanos.UTR
increased cell number | somatic clone, with Scer\GAL4NP5130
increased cell number | spermatogenesis, with Scer\GAL4VP16.nanos.UTR
increased mortality, with Scer\GAL4r4
increased occurrence of cell division | adult stage, with Scer\GAL4esg.PU
increased occurrence of cell division | adult stage, with Scer\GAL4NP0001
increased occurrence of cell division | adult stage, with Scer\GAL4NP5130
increased occurrence of cell division | adult stage, with Scer\GAL4NP7397
increased occurrence of cell division | adult stage | somatic clone, with Scer\GAL4NP5130
increased occurrence of cell division | embryonic stage | male, with Scer\GAL4VP16.nanos.UTR
increased occurrence of cell division | larval stage
increased size | adult stage
increased size | adult stage, with Scer\GAL4ey.PU
neoplasia | adult stage
neoplasia | spermatogenesis, with Scer\GAL4nanos.PU
some die during embryonic stage
some die during embryonic stage | recessive
some die during pupal stage
visible
visible, with Scer\GAL4bi-omb-Gal4
visible, with Scer\GAL4dpp.blk1
visible, with Scer\GAL4elav.PLu
visible, with Scer\GAL4ey.PH
visible, with Scer\GAL4GMR.PF
visible (with Df(1)os1A2)
visible (with Df(1)os1)
visible (with Df(1)oso)
visible | adult stage
visible | adult stage, with Scer\GAL4ey.PU
visible | pharate adult stage, with Scer\GAL4GMR.PU
visible | recessive
visible | somatic clone
Phenotype manifest in
Allele
abdominal segment 4
abdominal segment 5
abdominal segment 6
abdominal segment 7
abdominal segment 8
abdominal tergite
adult gut, with Scer\GAL4NP0001
adult midgut, with Scer\GAL4esg.PU
adult midgut, with Scer\GAL4NP5130
adult midgut epithelium, with Scer\GAL4NP7397, Scer\GAL4Tub.PU
adult midgut epithelium | adult stage, with Scer\GAL4NP5130
adult posterior midgut epithelium, with Scer\GAL4esg.PU
adult thoracic mechanosensory chaeta, with Scer\GAL4dpp.blk1
adult thoracic mechanosensory chaeta, with Scer\GAL4GMR15B03
adult thorax, with Scer\GAL4dpp.blk1
adult thorax, with Scer\GAL4GMR15B03
amnioserosa, with Scer\GAL4da.G32
border follicle cell
border follicle cell, with Scer\GAL4c306
border follicle cell, with Scer\GAL4slbo.2.6
border follicle cell, with Scer\GAL4upd1.PU
border follicle cell, with Scer\GAL4upd1-E132
border follicle cell (with upd1sisC-5)
border follicle cell (with upd1YM55)
border follicle cell | cell non-autonomous | ectopic | somatic clone, with Scer\GAL4Act5C.PP
border follicle cell | ectopic, with Scer\GAL4c306
border follicle cell | ectopic, with Scer\GAL4slbo.2.6
border follicle cell | ectopic | somatic clone, with Scer\GAL4Act5C.PI
border follicle cell | increased number, with Scer\GAL4c306
border follicle cell | increased number, with Scer\GAL4slbo.2.6
border follicle cell | somatic clone
centripetal follicle cell | ectopic | somatic clone, with Scer\GAL4Act5C.PP
cuticle & adult external head, with Scer\GAL4ey.PH
cyst cell of testis | increased number, with Scer\GAL4VP16.nanos.UTR
cyst progenitor cell, with Scer\GAL4VP16.nanos.UTR
cyst progenitor cell | ectopic | increased number, with Scer\GAL4VP16.nanos.UTR
cyst progenitor cell | increased number, with Scer\GAL4Act5C.PP
cyst progenitor cell | increased number, with Scer\GAL4VP16.nanos.UTR
ectoderm | extended germ band stage
egg chamber
embryonic/larval crystal cell | increased number | larval stage, with Scer\GAL4Hand.Δ
embryonic/larval crystal cell | increased number | larval stage, with Scer\GAL4NP1029
embryonic/larval cuticle
embryonic/larval gut | embryonic stage
embryonic/larval plasmatocyte
embryonic/larval proventriculus
embryonic abdominal segment 4
embryonic abdominal segment 5
embryonic abdominal segment 8
embryonic head, with Scer\GAL4h-1J3
embryonic hindgut, with Df(1)os1A2, Scer\GAL4byn-Gal4
embryonic hindgut, with Scer\GAL4byn-Gal4
embryonic metathoracic segment
enteroblast | adult stage | decreased number, with Scer\GAL4NP5130
escort cell | increased number, with Scer\GAL4C587
eye
eye, with Scer\GAL4bi-omb-Gal4
eye, with Scer\GAL4dpp.blk1
eye, with Scer\GAL4elav.PLu
eye, with Scer\GAL4ey.PH
eye, with Scer\GAL4ey.PU
eye, with Scer\GAL4GMR.PF
eye, with Scer\GAL4GMR.PU
eye (with Df(1)os1A2)
eye | dorsal, with Scer\GAL430A
eye | pharate adult stage, with Scer\GAL4GMR.PU
eye | somatic clone
eye | ventral, with Scer\GAL430A
eye-antennal disc
eye disc
eye disc, with Scer\GAL4GMR.PF
eye disc | cell non-autonomous | somatic clone | third instar larval stage, with Scer\GAL4Act5C.PI
eye disc | third instar larval stage, with Scer\GAL4dpp.blk1
eye disc | third instar larval stage, with Scer\GAL4ey.PH
eye disc anterior to the morphogenetic furrow, with Scer\GAL4GMR.PF
eye disc anterior to the morphogenetic furrow | larval stage
eye disc morphogenetic furrow, with Scer\GAL4ey.PH
female germline stem cell | increased number, with Scer\GAL4C587
filamentous actin | oogenesis, with Scer\GAL4upd1.PU
germarium, with Scer\GAL4C587
germline cell | embryonic stage, with Scer\GAL4VP16.nanos.UTR
germline cyst | increased number, with Scer\GAL4C587
germline stem cell, with Scer\GAL4VP16.nanos.UTR
germline stem cell | increased number, with Scer\GAL4VP16.nanos.UTR
gonad | ectopic & embryo
intestinal stem cell, with Scer\GAL4NP7397, Scer\GAL4Tub.PU
intestinal stem cell | adult stage, with Scer\GAL4NP7397
intestinal stem cell | adult stage | increased number, with Scer\GAL4NP5130
intestinal stem cell | increased number, with Scer\GAL4esg.PU
intestinal stem cell | somatic clone, with Scer\GAL4NP5130
intestinal stem cell of posterior adult midgut epithelium, with Scer\GAL4NP7397
larval head
main body follicle cell | somatic clone, with Scer\GAL4Act5C.PP
male germline cell | increased number, with Scer\GAL4A3.eya, Scer\GAL80ts.αTub84B
male germline stem cell | ectopic | increased number, with Scer\GAL4VP16.nanos.UTR
male germline stem cell | increased number, with Scer\GAL4Act5C.PP
male germline stem cell | increased number, with Scer\GAL4nanos.PG
male germline stem cell | increased number, with Scer\GAL4VP16.nanos.UTR
mesothoracic segment
mesothoracic tergum primordium | third instar larval stage, with Scer\GAL4ptc.PU
mitotic domain 1 | extended germ band stage, with Scer\GAL4da.G32
ommatidium
ommatidium, with Scer\GAL430A
ommatidium, with Scer\GAL4GMR.PF
ommatidium | increased number, with Scer\GAL4GMR.PF
ommatidium | somatic clone
ommatidium | somatic clone, with Scer\GAL4Act5C.PI
oocyte
oocyte, with Scer\GAL4upd1.PU
ovariole, with Scer\GAL4C587
perivitelline space, with Scer\GAL4upd1.PU
photoreceptor cell R8 | increased number, with Scer\GAL4GMR.PF
polar follicle cell
primary gonial cell | increased number, with Scer\GAL4Act5C.PP
primary gonial cell | increased number, with Scer\GAL4nanos.PG
primary gonial cell | increased number, with Scer\GAL4VP16.nanos.UTR
primordial germ cell | ectopic, with Scer\GAL4elav-C155
primordial germ cell | ectopic, with Scer\GAL4prd.RG1
primordial germ cell | ectopic, with Scer\GAL4twi.PG
round stem cell fusome | increased number, with Scer\GAL4Act5C.PP
round stem cell fusome | increased number, with Scer\GAL4VP16.nanos.UTR
somatic cell of testis | increased number, with Scer\GAL4A3.eya, Scer\GAL80ts.αTub84B
spermatogonium | increased number, with Scer\GAL4Act5C.PP
spermatogonium | increased number, with Scer\GAL4VP16.nanos.UTR
stalk follicle cell, with Scer\GAL4neur-P72
stalk follicle cell | increased number
stem cell fusome, with Scer\GAL4VP16.nanos.UTR
testis, with Scer\GAL4A3.eya, Scer\GAL80ts.αTub84B
testis, with Scer\GAL4nanos.PG
testis, with Scer\GAL4tj.PU
testis, with Scer\GAL4VP16.nanos.UTR
wing
wing, with Scer\GAL4Bx-MS1096
wing (with Df(1)os1)
wing (with Df(1)oso)
wing | posterior, with Scer\GAL4en-e16E
wing disc, with Scer\GAL4Bx-MS1096
wing disc, with Scer\GAL4ptc-559.1
wing pouch | third instar larval stage, with Scer\GAL4ptc.PU
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) (0)
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) (0)
Mus musculus (laboratory mouse) (0)
Xenopus tropicalis (Western clawed frog) (0)
Danio rerio (Zebrafish) (0)
Caenorhabditis elegans (Nematode, roundworm) (0)
Anopheles gambiae (African malaria mosquito) (0)
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:upd1. 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) (2)
upd2
2 of 13
upd3
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
upd1UAS.cUa
model of  cancer
CEA with Ras85DV12.UAS
(Wu et al., 2010)
upd1JF03149
model of  congenital leptin deficiency
CEA
(Beshel et al., 2017)
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 2 )
Allele
Disease
Interaction
References
upd1UAS.cZa
exacerbates  carcinoma
modeled by l(2)glGD4047
(Chatterjee et al., 2023)
upd1RNAi.UAS.cUa
ameliorates  cancer
modeled by Ras85DV12.UAS, l(2)glunspecified
(Wu et al., 2010)
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?
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
upd1
suppressible
BHD
(Singh et al., 2006)
upd1
suppressible
Brd
(Mukherjee et al., 2006)
upd1
suppressible
BRWD3
(Muller et al., 2005)
upd1
suppressible
BuGZ
(Wan et al., 2015)
upd1
suppressible
bun
(Mukherjee et al., 2006)
upd1
suppressible
CCT7
(Mukherjee et al., 2006)
upd1
suppressible
Cdk1
(Mukherjee et al., 2006)
upd1
suppressible
Cdk4
(Mukherjee et al., 2006)
upd1
suppressible
CG4306
(Mukherjee et al., 2006)
upd1
suppressible
CG5953
(Hu et al., 2025)
upd1
suppressible
CG17574
(Mukherjee et al., 2006)
upd1
suppressible
CG32982
(Mukherjee et al., 2006)
upd1
suppressible
Cip4
(Mukherjee et al., 2006)
upd1
suppressible
clu
(Mukherjee et al., 2006)
upd1
suppressible
cpo
(Mukherjee et al., 2006)
upd1
suppressible
crb
(Bach et al., 2003)
upd1
enhanceable
Csk
(Read et al., 2004)
upd1|egr
enhanceable
CtBP
(Worley et al., 2018)
upd1
enhanceable
CtBP
(Mukherjee et al., 2006, Bach et al., 2003)
upd1
suppressible
CtBP
(Mukherjee et al., 2006)
upd1
enhanceable
CycA
(Mukherjee et al., 2006)
upd1
enhanceable
CycD|Cdk4
(Chen et al., 2003)
upd1
suppressible
D
(Bach et al., 2003)
upd1
enhanceable
D
(Bach et al., 2003)
upd1
enhanceable
Dad
(Bach et al., 2003)
upd1
suppressible
dap
(Mukherjee et al., 2006)
upd1
suppressible
Delta
(Mukherjee et al., 2006)
upd1
suppressible
dind
(Zhao et al., 2022)
upd1
suppressible
dome
(Hombria et al., 2005, Chen et al., 2002, Ghiglione et al., 2002, Brown et al., 2001)
upd1
enhanceable
dome
(Bach et al., 2003)
Ras85D|upd1
suppressible
dome
(Wu et al., 2010)
upd1
suppressible
dpp
(Bach et al., 2003)
upd1
suppressible
E(bx)
(Badenhorst et al., 2002)
upd1
suppressible
Egfr
(Jiang et al., 2011)
upd1
suppressible
eIF3d1
(Wang et al., 2025)
upd1
suppressible
eIF4G1
(Wang et al., 2025)
upd1
suppressible
Elp6
(Mukherjee et al., 2006)
upd1
suppressible
esg
(Bach et al., 2003)
upd1
enhanceable
et
(Kallio et al., 2010)
upd1
suppressible
heca
(Bach et al., 2003)
upd1
suppressible
hh
(Bach et al., 2003)
upd1
enhanceable
Hipk
(Tettweiler et al., 2019)
upd1
suppressible
His2Av
(Bach et al., 2003)
upd1
suppressible
hop
(Bach et al., 2003, Chen et al., 2002, Kiger et al., 2001)
upd1
suppressible
Ipk2
(Seeds et al., 2015)
upd1
suppressible
jim
(Mukherjee et al., 2006)
upd1
enhanceable
jim
(Mukherjee et al., 2006)
upd1
suppressible
Krn
(Jiang et al., 2011)
upd1
suppressible
kto
(Bach et al., 2003)
upd1
suppressible
l(1)3Ag
(Bach et al., 2003)
upd1
suppressible
Lamp1
(Mukherjee et al., 2006)
upd1
suppressible
lilli
(Mukherjee et al., 2006)
upd1
suppressible
Mad
(Bach et al., 2003)
upd1
suppressible
mir-14
(Mukherjee et al., 2006)
upd1
suppressible
Mob2
(Mukherjee et al., 2006)
upd1
suppressible
mthl8
(Mukherjee et al., 2006)
upd1
suppressible
Myc
(Mukherjee et al., 2006)
upd1
suppressible
N
(Mukherjee et al., 2006)
upd1
enhanceable
NFAT
(Mukherjee et al., 2006)
upd1
suppressible
NFAT
(Mukherjee et al., 2006)
upd1
suppressible
pAbp
(Mukherjee et al., 2006)
upd1
suppressible
Phb1
(Zhao et al., 2022)
upd1
suppressible
Phb2
(Zhao et al., 2022)
upd1
suppressible
pie
(Bach et al., 2003)
upd1
suppressible
put
(Bach et al., 2003)
upd1
suppressible
px
(Bach et al., 2003)
upd1
suppressible
Ras85D
(Jiang et al., 2011)
upd1
suppressible
Rho1
(Bertet et al., 2009)
upd1
suppressible
rux
(Mukherjee et al., 2006)
upd1
suppressible
sax
(Bach et al., 2003)
upd1
suppressible
shg
(Mallart et al., 2024)
upd1
enhanceable
Socs36E
(Bach et al., 2003)
upd1
suppressible
Socs36E
(Singh et al., 2010)
upd1
suppressible
Sord2
(Mukherjee et al., 2006)
upd1
suppressible
Sox100B
(Jin et al., 2020)
upd1
enhanceable
spri
(Mukherjee et al., 2006)
upd1
suppressible
spri
(Mukherjee et al., 2006)
upd1
suppressible
sqh
(Bertet et al., 2009)
upd1
suppressible
Ssdp
(Mukherjee et al., 2006)
upd1
suppressible
Stat92E
(Read et al., 2004)
upd1
enhanceable
Ten-m
(Mukherjee et al., 2006)
upd1
suppressible
tkv
(Bach et al., 2003)
upd1
suppressible
trbl
(Mukherjee et al., 2006)
upd1
suppressible
upd2
(Beshel et al., 2017)
upd1|Ras85D
enhanceable
upd2
(Wu et al., 2010)
Ras85D|l(2)gl|upd1
suppressible
upd2
(Wu et al., 2010)
upd1
suppressible
WASp
(Bertet et al., 2009)
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
Alms1a
suppressible
upd1
(Chen and Yamashita, 2020)
bi
suppressible
upd1
(Tsai et al., 2015)
Bub3
suppressible
upd3|upd1
(Romão et al., 2021)
Delta
suppressible
upd1
(Reynolds-Kenneally and Mlodzik, 2005)
drm
suppressible
upd1
(Johansen et al., 2003)
dysf
suppressible
upd1
(Wu et al., 2022)
Egfr
suppressible
upd1
(Herranz et al., 2012)
egr|CtBP
enhanceable
upd1
(Worley et al., 2018)
eyg
suppressible
upd1
(Chao et al., 2004)
flw
enhanceable
upd1
(Sun et al., 2011)
gcm
suppressible
upd1
(Jacques et al., 2009)
hid
suppressible
upd1
(Herz et al., 2006)
hop
enhanceable
upd1
(McGregor et al., 2002, Harrison et al., 1998)
hop
suppressible
upd1
(Zhao et al., 2022)
L
enhanceable
upd1
(Wang and Huang, 2009)
mir-279
suppressible
upd1
(Luo and Sehgal, 2012)
N
suppressible
upd1
(Djiane et al., 2013)
Rac1
suppressible
upd1
(Silver, 2005)
Ras85D
enhanceable
upd1
(Willecke et al., 2011)
Ras85D|l(2)gl
suppressible
upd1
(Wu et al., 2010)
rpr
enhanceable
upd1
(Harris et al., 2020)
Ser
suppressible
upd1
(Reynolds-Kenneally and Mlodzik, 2005)
External Data
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.
SignaLink - A signaling pathway resource with multi-layered regulatory networks.
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
X
Recombination map
1-60
Cytogenetic map
17A5-17A5
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
17A5-17A5
Limits computationally determined from genome sequence between P{EP}ari-1EP317 and P{EP}EP1378
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
17A5-17A7
(Sefton, 2000.2.4)
17A-17A
(Harrison, 1998.6.17)
17A-17A
(determined by in situ hybridisation)
(Cline, 1993)
17A5-17A8
(determined by in situ hybridisation)
(Eberl et al., 1992)
Experimentally Determined Recombination Data
Location

1-60

(FlyBase, 2016)

1-59.2

(Comeron et al., 2012, University of Buenos Aires, 1951, Bridges and Brehme, 1944)

1-58.7

(Cline, 1993)
Left of (cM)
Right of (cM)
Notes

Mapping based on oso.

(Bridges and Brehme, 1944)
Stocks and Reagents
Stocks (33)
Genomic Clones (12)
 

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

cDNA Clones (4)
 

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

    (Beshel et al., 2017)

    monoclonal

    (Zhang et al., 2013)
    Commercially Available Antibodies
     
    Cell Line Information
    Publicly Available Cell Lines
     
      Other Stable Cell Lines
       
      Other Comments

      upd1 is required to maintain basal turnover of the midgut epithelium by controlling intestinal stem cell maintenance in an autocrine manner.

      (Osman et al., 2012)

      Gene expression is increased in response to the presence of either one or two copies of Scer\GAL4hs.PB.

      (Liu and Lehmann, 2008)

      upd1 protein, secreted by the polar follicle cells, may act as a morphogen to specify anterior-posterior pattern in the ovarian follicle cells.

      (Xi et al., 2003)

      upd1 is required for self-renewal of male germ line stem cells and somatic cyst progenitor cells.

      (Tulina and Matunis, 2001)

      upd1 acts as an X-linked signal element in sex determination.

      (Sefton et al., 2000)

      A localised source of upd1 protein, which is present at the midline of the developing eye, is capable of activating the JAK/STAT pathway over long distances.

      (Zeidler et al., 1999)

      upd1 is necessary to determine the position of the eye equator via regulation of mirr.

      (Zeidler et al., 1999)

      γ ray induced loss of function mutations suppress male-lethal duplications of numerator elements.

      (Cline, 1993)

      The os gene has been studied as part of a genetic and developmental study of polytene section 17 of the X chromosome. A duplication of this region, fcl+Y, has been constructed to allow mutagenesis screens and complementation analyses of the region.

      (Eberl et al., 1992)

      upd1 mutants display defects predominantly in mesothorax and fifth row, head defects and eighth segment defects.

      (Wieschaus et al., 1984)
      Relationship to Other Genes
      Source for database merge of

      Source for merge of: os sisC

      (Sefton et al., 1998)
      Additional comments

      Classical alleles of the 'os' (outstretched) complementation group affect expression of both the upd1 and upd3 genes and map to untranscribed regions, suggesting that they disrupt regulatory elements shared by both genes.

      (Wang et al., 2014)

      "upd" and "os" can be genetically separated. This raises the possibility that "os" function may reside with an "upd-like" gene ("upd2" or "upd3").

      (Wang, 2005)
      Nomenclature History
      Source for database identify of

      Source for identity of: os CG5993

      (Li, 1999.11)
      Nomenclature comments
      Etymology
      Synonyms and Secondary IDs (34)
      Reported As
      Symbol Synonym
      CG5993
      (Meadows, 2017.9.14, Wright et al., 2011, Ni et al., 2010.12.1, Perkins et al., 2009, Grieder et al., 2007, Klebes et al., 2005)
      OS
      (Mbodj et al., 2013, Arbouzova and Zeidler, 2006)
      Os
      (Lin et al., 2015, Salazar-Jaramillo et al., 2014, Payne and Braun, 2006, Terry et al., 2006)
      UPD
      (He et al., 2025, Starz-Gaiano et al., 2009, Arbouzova and Zeidler, 2006, Castelli-Gair Hombria, 2006, Scadden, 2006, Silver and Montell, 2002)
      UPD1
      (Mahishi et al., 2024, Toprak et al., 2020)
      Unpaireds
      (Jasper, 2020)
      Upd
      (Li et al., 2025, Mpamhanga and Kounatidis, 2024, Chatterjee et al., 2023, Kietz and Meinander, 2023, Li et al., 2023, Stączek et al., 2023, Zhai et al., 2023, Enomoto and Igaki, 2022, Watase et al., 2022, Zhao et al., 2022, Casares and McGregor, 2021, Finger et al., 2021, Lam Wong and Verheyen, 2021, Rosendo Machado et al., 2021, Surkova et al., 2021, Vidaurre and Chen, 2021, Antel and Inaba, 2020, Hao et al., 2020, Hayashi et al., 2020, Horne-Badovinac, 2020, Merkle et al., 2020, Morata and Calleja, 2020, Peercy and Starz-Gaiano, 2020, Zhang and Cai, 2020, Angulo et al., 2019, Kim et al., 2019, Loyola et al., 2019, Powers and Srivastava, 2019, Ahmed-de-Prado and Baonza, 2018, Enomoto et al., 2018, Mehrotra and Deshpande, 2018, Perochon et al., 2018, Petrovsky et al., 2018, Richardson and Portela, 2018, Stuelten et al., 2018, Wu et al., 2018, Feng et al., 2017, Kashio et al., 2017, Khanna et al., 2017, Kim et al., 2017, Tamori and Deng, 2017, Deshpande et al., 2016, Fisher et al., 2016, Jiang et al., 2016, Mbodj et al., 2016, Yadav et al., 2016, Dorn and Dorn, 2015, Enomoto et al., 2015, Greenspan et al., 2015, Merkling et al., 2015, Nie et al., 2015, Seeds et al., 2015, Thomas et al., 2015, Wang et al., 2015, Chabu and Xu, 2014, Chen et al., 2014, Huang et al., 2014, Tamori and Deng, 2014, Ayyaz and Jasper, 2013, Bausek, 2013, Hombría and Sotillos, 2013, Li et al., 2013, Morin-Poulard et al., 2013, Pastor-Pareja and Xu, 2013, Ren et al., 2013, Sato et al., 2013, Shim et al., 2013, Southall et al., 2013, Takashima et al., 2013, Yamashita, 2013, Zeidler and Bausek, 2013, Zhou et al., 2013, Zoranovic et al., 2013, Amoyel and Bach, 2012, Grönholm et al., 2012, Hayashi et al., 2012, Kounatidis and Ligoxygakis, 2012, Lim et al., 2012, Luo and Sehgal, 2012, Matunis et al., 2012, Monk et al., 2012, White-Cooper, 2012, Zoller and Schulz, 2012, Apidianakis and Rahme, 2011, Buechling et al., 2011, Jiang et al., 2011, Stec and Zeidler, 2011, Van De Bor et al., 2011, Willecke et al., 2011, Wright et al., 2011, Yoon et al., 2011, Bina et al., 2010, Lin et al., 2010, Ren et al., 2010, Akbar et al., 2009, Gilbert et al., 2009, González et al., 2009, Jiang et al., 2009, Sheng et al., 2009, Vaccari et al., 2009, Kim et al., 2008, Rivas et al., 2008, Yamashita and Fuller, 2008, Fuller and Spradling, 2007, Luque and Milan, 2007, Nurminsky, 2007, Pei and Harrison, 2007, Tolwinski, 2007, Tsai et al., 2007, Yasugi et al., 2007, Bartscherer et al., 2006, Bina and Zeidler, 2006, Giebel and Wodarz, 2006, Sexton et al., 2006, Wong et al., 2006, Baeg et al., 2005, Moberg et al., 2005, Muller et al., 2005, Reynolds-Kenneally and Mlodzik, 2005, Thompson et al., 2005, Vaccari and Bilder, 2005, Yamashita et al., 2005, Zeidler et al., 2005, Zhang and Li, 2005, Ekas and Bach, 2004, Fuchs et al., 2004, Gilboa and Lehmann, 2004, Oh et al., 2004, Read et al., 2004, Sexton et al., 2004, Agaisse et al., 2003, Arbouzova et al., 2003, Denef and Schupbach, 2003, Hultmark and Ekengren, 2003, Mukherjee and Zeidler, 2003, Wallenfang and DiNardo, 2003, Xi et al., 2003, Kisseleva et al., 2002, Tulina and Matunis, 2002, Xi and Harrison, 2002, Tulina and Matunis, 2001, Blair, 1999, Zeidler et al., 1999, Zeidler et al., 1999)
      Upd1
      (Parvez and Rahul, 2026, Wang et al., 2026, Wu et al., 2026, Diaz et al., 2025, Djiane, 2025, Hirooka et al., 2025, Krejčová and Bajgar, 2025, Park et al., 2025, Hersperger et al., 2024, Tran et al., 2024, Wang et al., 2024, Neophytou and Pitsouli, 2022, Bilder et al., 2021, Dillard et al., 2021, Gillette et al., 2021, Harnish et al., 2021, Järvelä-Stölting et al., 2021, Kim et al., 2021, Lee et al., 2021, Schneider and Imler, 2021, King and Sehgal, 2020, Koyama et al., 2020, Krautz et al., 2020, Wang and Spradling, 2020, Zhou et al., 2020, Nässel, 2018, Daisley et al., 2017, Gervais and Bardin, 2017, Liu and Jin, 2017, Subramanian et al., 2017, Vanha-Aho et al., 2016, Guo et al., 2014, Wang et al., 2014, Morin-Poulard et al., 2013, Liu et al., 2012, Apidianakis and Rahme, 2011, Biteau et al., 2011, Zhao et al., 2011, Devergne et al., 2007)
      l(1)YC43
      l(1)YM55
      od
      (University of Buenos Aires, 1951)
      odsy
      os
      (Tettweiler et al., 2019, Atkins et al., 2016, Zhang et al., 2015, Chen et al., 2013, Toledano et al., 2012, Kaplan et al., 2011, Nien et al., 2011, Zheng et al., 2011, Ekas et al., 2010, Popodi et al., 2010-, Venken et al., 2010, Almudi et al., 2009, Bertet et al., 2009, Gutierrez-Aviño et al., 2009, Jacques et al., 2009, Wang and Huang, 2009, Liu and Lehmann, 2008, Wang and Harrison, 2008, Grieder et al., 2007, Sturgill et al., 2007, Wang and Harrison, 2007, Manak et al., 2006, Mukherjee et al., 2006, Wang et al., 2006, Möller et al., 2005, Chao et al., 2004, Sorrentino et al., 2004, Tsai and Sun, 2004, Agaisse et al., 2003, Betz et al., 2001, Tsai and Sun, 2001, Harrison et al., 1998, Lindsley and Zimm, 1992, Verderosa and Muller, 1954)
      sis-C
      (Fisher and Caudy, 1998)
      sis-c
      (Calhoun et al., 2004, Pettus and Nagoshi, 1998)
      sisC
      (Nien et al., 2011, Liang et al., 2008, Avila and Erickson, 2007, Pomiankowski et al., 2004, Schutt and Nothiger, 2000, Sefton et al., 2000, Zeidler et al., 2000, Sefton et al., 1998, Cline, 1993)
      sy
      (Patterson, 1932)
      unp
      (Badenhorst et al., 2002)
      upa
      (Martinez Arias, 1993)
      upd
      (Urban et al., 2026, Zheng et al., 2026, Nelson et al., 2025, Eslahi et al., 2024, Wen et al., 2024, Chandrasekhara et al., 2023, Kanaoka et al., 2023, Ku et al., 2023, Cao et al., 2022, Evans et al., 2022, Hayashi et al., 2022, Mallart et al., 2022, Gong et al., 2021, Herrera and Bach, 2021, Chen and Yamashita, 2020, Chen et al., 2020, Harsh et al., 2020, Jin et al., 2020, Moore et al., 2020, Xu et al., 2020, Sreejith et al., 2019, Wittes and Schüpbach, 2019, Baker et al., 2018, Chen et al., 2018, Crossman et al., 2018, Fisher et al., 2018, Palmer et al., 2018, Parvy et al., 2018, Petrovsky and Großhans, 2018, Saadin and Starz-Gaiano, 2018, Recasens-Alvarez et al., 2017, Takemura and Nakato, 2017, Torres et al., 2017, Vollmer et al., 2017, Zhang et al., 2017, Fregoso Lomas et al., 2016, Saadin and Starz-Gaiano, 2016, Shen et al., 2016, Singh et al., 2016, Barr et al., 2015, Das and Dobens, 2015, Glassford et al., 2015, Katsuyama et al., 2015, Matsuda et al., 2015, Pasco et al., 2015, Pinto et al., 2015, Ren et al., 2015, Santabárbara-Ruiz et al., 2015, Singh, 2015, Sun et al., 2015, Tataroglu and Emery, 2015, Tsai et al., 2015, Wan et al., 2015, Zang et al., 2015, Li et al., 2014, Lin et al., 2014, Okumura et al., 2014, Scopelliti et al., 2014, Shields et al., 2014, Sopko et al., 2014, Voog et al., 2014, Wang et al., 2014, Xu and Cherry, 2014, Aleksic et al., 2013, Borensztejn et al., 2013, Geisbrecht et al., 2013, Guo et al., 2013, Hombría and Serras, 2013, Hombría and Sotillos, 2013, Katsuyama et al., 2013, Kemp et al., 2013, Li et al., 2013, Lin et al., 2013, Lucas et al., 2013, Morillo Prado et al., 2013, Sinha et al., 2013, Wang et al., 2013, Wells et al., 2013, Yadlapalli and Yamashita, 2013, Zeidler and Bausek, 2013, Zhang et al., 2013, Zoranovic et al., 2013, Herranz et al., 2012, Issigonis and Matunis, 2012, Matunis et al., 2012, Toledano et al., 2012, Toledano et al., 2012, Tran et al., 2012, Wong and Jones, 2012, Casper et al., 2011, Copf et al., 2011, de Cuevas and Matunis, 2011, Feng et al., 2011, Johnson et al., 2011, Kim et al., 2011, Morris and Spradling, 2011, Parrott et al., 2011, Sinenko et al., 2011, Singh et al., 2011, Stec and Zeidler, 2011, Sun et al., 2011, Uddin et al., 2011, Wang et al., 2011, Wang et al., 2011, Wright et al., 2011, Yuva-Aydemir et al., 2011, Zheng et al., 2011, Beebe et al., 2010, Flaherty et al., 2010, Gladstein et al., 2010, Goto et al., 2010, Kallio et al., 2010, Liu et al., 2010, Martinez and Cavalli, 2010, Ngo et al., 2010, Shaw et al., 2010, Singh et al., 2010, Sotillos et al., 2010, Wu et al., 2010, Zhai et al., 2010, Almudi et al., 2009, Bertet et al., 2009, Casper and Van Doren, 2009, Classen et al., 2009, Cronin et al., 2009, Flaherty et al., 2009, Gutierrez-Aviño et al., 2009, Papagiannouli and Mechler, 2009, Sheng et al., 2009, Smith-Bolton et al., 2009, Wang and Huang, 2009, Cinnamon et al., 2008, Copf and Preat, 2008, Duong et al., 2008, Leatherman and DiNardo, 2008, López-Onieva et al., 2008, Pastor-Pareja et al., 2008, Sexton and Harrison, 2008, Sotillos et al., 2008, Voog et al., 2008, Wang and Harrison, 2008, Wang et al., 2008, Wang et al., 2008, Yasugi et al., 2008, Assa-Kunik et al., 2007, Avila and Erickson, 2007, Ayala-Camargo et al., 2007, Bach et al., 2007, Boyle et al., 2007, Devergne et al., 2007, Flaherty et al., 2007, Furriols et al., 2007, Grieder et al., 2007, Sexton and Harrison, 2007, Singh et al., 2007, Tsai et al., 2007, Tsai et al., 2007, Wang and Harrison, 2007, Wang et al., 2007, Arbouzova et al., 2006, Brown et al., 2006, DEVERGNE and NOSELLI, 2006, Ekas et al., 2006, Ekas et al., 2006, Herz, 2006, Herz et al., 2006, Keller, 2006, Lovegrove et al., 2006, Lovegrove et al., 2006, Montell, 2006, Mukherjee et al., 2006, Singh et al., 2006, Singh et al., 2006, Wang et al., 2006, Wang et al., 2006, Zheng et al., 2006, Davies et al., 2005, Decotto and Spradling, 2005, Ekas and Bach, 2005, Gilbert et al., 2005, Grieder et al., 2005, Hombria et al., 2005, Liu and Sun, 2005, Mukherjee et al., 2005, Muller, 2005, Reynolds-Kenneally and Mlodzik, 2005, Sexton et al., 2005, Tsai and Sun, 2005, Wang, 2005, Wang et al., 2005, Wawersik et al., 2005, Xie et al., 2005, Agaisse and Perrimon, 2004, Josten et al., 2004, Rawlings et al., 2004, Starz-Gaiano and Montell, 2004, Tsai and Sun, 2004, Tsai and Sun, 2004, Yamashita et al., 2004, Bach et al., 2003, Brown et al., 2003, Chen et al., 2003, Johansen et al., 2003, Johansen et al., 2003, Johansen et al., 2003, Li et al., 2003, Maines et al., 2003, Tulina et al., 2003, Xi et al., 2003, Adler, 2002, Bach and Perrimon, 2002, Bach and Perrimon, 2002, Badenhorst et al., 2002, Beccari et al., 2002, Castelli-Gair Hombria and Brown, 2002, Chen et al., 2002, Chen et al., 2002, Ghiglione et al., 2002, Grammont and Irvine, 2002, Green et al., 2002, Harrison et al., 2002, Harrison et al., 2002, Hou et al., 2002, Johansen et al., 2002, Karsten et al., 2002, Lengyel and Iwaki, 2002, Lin, 2002, McGregor et al., 2002, Zhao and Garbers, 2002, Zheng and Hou, 2002, Brown et al., 2001, Iwaki et al., 2001, Johansen et al., 2001, Johansen et al., 2001, Jones et al., 2001, Kiger et al., 2001, Luo and Dearolf, 2001, Silver and Montell, 2001, Spradling et al., 2001, Tsai and Sun, 2001, Xi and Harrison, 2001, Bach et al., 2000, Jinks et al., 2000, Morey et al., 2000, Xi and Harrison, 2000, Zeidler et al., 2000, Bach et al., 1999, Dearolf, 1999, Keating and Harrison, 1999, Morey and Harrison, 1999, Savant-Bhonsale et al., 1999, Tsai and Sun, 1999, Zeidler et al., 1999, Zeidler et al., 1999, Harrison et al., 1998, Mathey-Prevot et al., 1998, Morey and Harrison, 1998, Sefton et al., 1998, Harrison and Perrimon, 1997, McCoon and Gilman, 1997, Sefton et al., 1997, Cline and Meyer, 1996, Harbecke and Lengyel, 1995, Klingler and Gergen, 1993, Parkhurst and Ish-Horowicz, 1991, Perrimon et al., 1989, DiNardo and O'Farrell, 1987, Carroll and Scott, 1986, Wieschaus et al., 1984)
      upd1
      (Cruz et al., 2026, Strand, 2026, Cong et al., 2025, Dong et al., 2025, Geng et al., 2025, He et al., 2025, Kong et al., 2025, Kumar et al., 2025, Sun et al., 2025, Balakireva et al., 2024, Guo et al., 2024, Ju et al., 2024, Li et al., 2024, Li et al., 2024, Liu et al., 2024, Miao et al., 2024, Pfefferkorn et al., 2024, Barrio et al., 2023, Chen et al., 2023, Floc'hlay et al., 2023, Jiang et al., 2023, Molina-Gil et al., 2023, Neophytou et al., 2023, Xu et al., 2023, Yamada et al., 2023, Zhou and Boutros, 2023, Deshpande et al., 2022, Gera et al., 2022, Shen et al., 2022, Weichselberger et al., 2022, Worley and Hariharan, 2022, Destalminil-Letourneau et al., 2021, Dong et al., 2021, Fabian et al., 2021, Gogia et al., 2021, Joy et al., 2021, Maurya et al., 2021, Sanhueza et al., 2021, Slaidina et al., 2021, Takemura et al., 2021, Harris et al., 2020, La Marca and Richardson, 2020, Maselko et al., 2020, Rust et al., 2020, Statzer and Ewald, 2020, Vizcaya-Molina et al., 2020, Wei et al., 2020, Asri et al., 2019, Banerjee et al., 2019, Camara et al., 2019, Gultekin and Steller, 2019, Herrera and Bach, 2019, Hudry et al., 2019, Khezri and Rusten, 2019, Kwasnieski et al., 2019, Lin et al., 2019, Lin et al., 2019, Moskalev et al., 2019, Nelson et al., 2019, Reedy et al., 2019, Saavedra and Perrimon, 2019, Tudrej et al., 2019, Xu et al., 2019, Yang et al., 2019, Jia et al., 2018, Prange et al., 2018, Sayin et al., 2018, Beshel et al., 2017, Katheder et al., 2017, Lee et al., 2017, Péan et al., 2017, Transgenic RNAi Project members, 2017-, Xu et al., 2017, Hoi et al., 2016, Shih et al., 2016, Toggweiler et al., 2016, Ayyaz et al., 2015, Doggett et al., 2015, Woodcock et al., 2015, Zhai et al., 2015, Djiane et al., 2013, Osman et al., 2012, Osman et al., 2012, Rajan and Perrimon, 2012, Krzemień et al., 2007)
      Name Synonyms
      Outstretched
      (Zoranovic et al., 2013, Amoyel and Bach, 2012, Matunis et al., 2012, Leatherman and DiNardo, 2007)
      Unpaired
      (St Johnston, 2023, Milas and Telley, 2022, Hayashi et al., 2020, Zhang and Cai, 2020, Khanna et al., 2017, Tamori and Deng, 2017, Wang et al., 2014, Bausek, 2013, Morin-Poulard et al., 2013, Wang et al., 2013, Wells et al., 2013, Yamashita, 2013, Zoranovic et al., 2013, Amoyel and Bach, 2012, Lim et al., 2012, Luo and Sehgal, 2012, Matunis et al., 2012, Toledano et al., 2012, Zoller and Schulz, 2012, Yoon et al., 2011, Bina et al., 2010, Classen et al., 2009, Smith-Bolton et al., 2009, Vaccari et al., 2009, Sotillos et al., 2008, Yasugi et al., 2008, Ayala et al., 2007, Fuller and Spradling, 2007, Luque and Milan, 2007, Tountas and Fortini, 2007, Yasugi et al., 2007, Ekas et al., 2006, Montell, 2006, Morrison and Kimble, 2006, Scadden, 2006, Seydoux and Braun, 2006, Lopez-Schier, 2003, Solnica-Krezel and Eaton, 2003, Wallenfang and DiNardo, 2003, Rorth, 2002, Morey et al., 2001, Wasserman and DiNardo, 2001, Harrison and Morey, 2000)
      Unpaired 1
      (Devergne et al., 2007)
      Unpaired/Out-stretched
      (Payne and Braun, 2006)
      outstretched
      (Piccirillo et al., 2014, Ekas et al., 2010, Hill-Burns and Clark, 2009, Kurusu et al., 2008, Musters et al., 2006, University of Buenos Aires, 1951)
      outstretched small eye
      (Lindsley and Zimm, 1992)
      outstretched-smalleye
      sisterless c
      (Cline, 1993)
      small-eye
      (Morgan, 1938)
      unpaired
      (Chan et al., 2023, Palmer et al., 2018, Kashio et al., 2017, Wieschaus and Nüsslein-Volhard, 2016, Matsuda et al., 2015, Okumura et al., 2014, Guo et al., 2013, Katsuyama et al., 2013, Lucas et al., 2013, Sinha et al., 2013, Zhang et al., 2013, Grönholm et al., 2012, Kim et al., 2011, Singh et al., 2011, Uddin et al., 2011, Wang et al., 2011, Beebe et al., 2010, Kallio et al., 2010, Zhai et al., 2010, Bertet et al., 2009, Cronin et al., 2009, Gutierrez-Aviño et al., 2009, Lin, 2008, Wang et al., 2008, Avila and Erickson, 2007, Ayala-Camargo et al., 2007, Bach et al., 2007, Boyle et al., 2007, Singh et al., 2007, Tsai et al., 2007, Tsai et al., 2007, Casper and Van Doren, 2006, Herz et al., 2006, Maurange et al., 2006, Singh et al., 2006, Agaisse and Perrimon, 2004, Starz-Gaiano and Montell, 2004, Tsai and Sun, 2004, Wawersik et al., 2004, Torres et al., 2003, Kile et al., 2002, Rennebeck et al., 2002, Johansen et al., 2001, Matunis and Tulina, 2001, Matunis and Tulina, 2001, Rennebeck and Harrison, 2001, Lemaitre, 2000.12.20, Williams, 2000, Xi and Harrison, 2000, Zeidler and Perrimon, 2000, Blair, 1999, Mlodzik, 1999, Strutt and Strutt, 1999, Harrison, 1998.6.17, Harrison et al., 1998, Morey and Harrison, 1998, Hou and Perrimon, 1997, Martinez Arias, 1993, Carroll and Scott, 1986)
      unpaired 1
      (Kula-Eversole and Hatini, 2007)
      unpaired1
      (Shi et al., 2020)
      upd homolog
      (Feng et al., 2012)
      Secondary FlyBase IDs
      • FBgn0003012
      • FBgn0003956
      • FBgn0016985
      Datasets (0)
      Study focus (0)
      Experimental Role
      Project
      Project Type
      Title
      Study result (0)
      Result
      Result Type
      Title
      External Crossreferences and Linkouts ( 49 )
      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/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
      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.
      SignaLink - A signaling pathway resource with multi-layered regulatory networks.
      References (753)