FlyBase Gene Report: Dmel\upd2

General Information

Symbol

Dmel\upd2

Species

D. melanogaster

Name

unpaired 2

Annotation Symbol

CG5988

Feature Type

protein_coding_gene

FlyBase ID

FBgn0030904

Gene Model Status

Current

Stock Availability

12 publicly available

Gene Summary

unpaired 2 (upd2) encodes a secreted molecule that acts at a distance as a ligand for the JAK/STAT signal transduction pathway. upd2 mutants are viable due to redundancy with other Upd-family genes. [Date last reviewed: 2019-03-21] (FlyBase Gene Snapshot)

All Summaries

Gene Snapshot
Alliance
Auto summary
Pathway
Gene Group
Interactive Fly

Also Known As

Upd, upd-2

Key Links

Genomic Location

Cytogenetic map

17A3-17A3

Sequence location

X:18,240,654..18,243,284 [-]

Recombination map

1-60

RefSeq locus

NC_004354 REGION:18240654..18243284

Sequence

Get Decorated FASTA

Genomic Maps

JBrowse

Other Genome Views

The following external sites may use different assemblies or annotations than FlyBase.

Function

Gene Ontology (GO) Annotations (12 terms)

Molecular Function (1 term)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

enables cytokine activity

inferred from genetic interaction with FLYBASE:dome; FB:FBgn0043903

(Hombria et al., 2005)

inferred from direct assay

(Moore et al., 2020, Wright et al., 2011)

Terms Based on Predictions or Assertions (0 terms)

Biological Process (9 terms)

Terms Based on Experimental Evidence (8 terms)

CV Term

Evidence

References

involved_in cell surface receptor signaling pathway via JAK-STAT

inferred from genetic interaction with FLYBASE:hop; FB:FBgn0004864

(Bailetti et al., 2019, Wright et al., 2011)

inferred from mutant phenotype

(Osman et al., 2012, Rajan and Perrimon, 2012, Wawersik et al., 2005)

inferred from direct assay

(Moore et al., 2020, Wright et al., 2011, Vidal et al., 2010, Hombria et al., 2005)

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 intestinal epithelial structure maintenance

inferred from mutant phenotype

(Osman et al., 2012)

involved_in lamellocyte differentiation

inferred from mutant phenotype

(McMullen et al., 2024)

involved_in paracrine signaling

inferred from direct assay

(Wright et al., 2011)

involved_in positive regulation of lipid storage

inferred from mutant phenotype

(Rajan et al., 2017)

involved_in positive regulation of multicellular organism growth

inferred from mutant phenotype

(Rajan and Perrimon, 2012)

involved_in segment specification

inferred from mutant phenotype

(Hombria et al., 2005)

involved_in tissue regeneration

inferred from mutant phenotype

(Harris et al., 2020)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

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-)

Cellular Component (2 terms)

Terms Based on Experimental Evidence (2 terms)

CV Term

Evidence

References

located_in extracellular region

inferred from direct assay

(Wright et al., 2011)

is_active_in extracellular space

inferred from direct assay

(Vidal et al., 2010)

located_in extracellular space

inferred from direct assay

(Rajan et al., 2017, Hombria et al., 2005)

Terms Based on Predictions or Assertions (0 terms)

Gene Group (FlyBase)

UNPAIRED FAMILY

Pathway (FlyBase)

JAK-STAT SIGNALING PATHWAY CORE COMPONENTS

Protein Family (UniProt)

Protein Signatures (InterPro)

Unpaired protein

Summaries

Gene Snapshot

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).

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

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\upd2 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 Q9VWX2)

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 5.53

Gene model reviewed during 6.23

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

upd2-RC

FBtr0473620

NM_001370039

1980

406

Additional Transcript Data and Comments

Reported size (kB)

Comments

External Data

Crossreferences

Polypeptide Data

Annotated Polypeptides

Name

FlyBase ID

Predicted MW (kDa)

Length (aa)

Theoretical pI

UniProt

RefSeq ID

GenBank

upd2-PC

FBpp0423083

45.6

406

9.75

AAF48815

Polypeptides with Identical Sequences

There is only one protein coding transcript and one polypeptide associated with this gene

Additional Polypeptide Data and Comments

Reported size (kDa)

Comments

External Data

Crossreferences

InterPro - A database of protein families, domains and functional sites

Unpaired protein

Linkouts

Sequences Consistent with the Gene Model

Nucleotide / Polypeptide Records

UniProt

Q9VWX2

Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\upd2 using the Feature Mapper tool.

External Data

Crossreferences

Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.

upd2_1

Linkouts

Expression Data

Testis-specificity index

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

-0.19

Transcript Expression

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 9

organism | segmentally repeated | striped

(Hombria et al., 2005)

embryonic stage 10

tracheal section

Comment: tracheal expression is transient

(Hombria et al., 2005)

embryonic stage 11

hindgut

(Hombria et al., 2005)

posterior spiracle primordium

(Hombria et al., 2005)

Additional Descriptive Data

upd2 is expressed in segmentally repeated stripes at embryonic stage 9. At embryonic stage 10, upd2 has transient tracheal expression. At embryonic stage 11, the tracheal expression of upd2 has been replaced with expression in the hindgut and posterior spiracles.

(Hombria et al., 2005)

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in extracellular region

inferred from direct assay

(Wright et al., 2011)

is_active_in extracellular space

inferred from direct assay

(Vidal et al., 2010)

located_in extracellular space

inferred from direct assay

(Rajan et al., 2017, Hombria et al., 2005)

Expression Deduced from Reporters

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\upd2 in JBrowse

RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region

View exonic expression by developmental stage for Dmel\upd2

View exonic expression by tissue for Dmel\upd2

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

32805/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0030904

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0030904

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0030904

Flygut - An atlas of the Drosophila adult midgut

FBgn0030904

Images

FlyBase Wiki

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 7 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 24 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of upd2

Transgenic constructs containing regulatory region of upd2

Aberrations (Deficiencies and Duplications) ( 2 )

Inferred from experimentation ( 2 )

Gene disrupted in

Df(1)os1A2

(Hombria et al., 2005)

Gene not disrupted in

Df(1)os^UE69

(Hombria et al., 2005)

Inferred from location ( 14 )

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 | female, with Spyo\Cas9^{dxCas9-3.7::VPR.αTub84B}

upd2^TOE.GS00171

lethal - all die before end of P-stage, with Spyo\Cas9^{dCas9::VPR.foxo}

upd2^TOE.GS00171

lethal - all die before end of P-stage, with Spyo\Cas9^{dCas9::VPR.wg-ΔC1}

upd2^TOE.GS00171

lethal - all die before end of P-stage, with Spyo\Cas9^{dCas9::VPR.αTub84B}

upd2^TOE.GS00171

viable

upd2^Δ

viable, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

upd2^GD6513

viable, with Scer\GAL4^Mef2.PR

upd2^GD6513

viable, with Scer\GAL4^pnr-MD237

upd2^GD6513

viable | male, with Spyo\Cas9^{dxCas9-3.7::VPR.αTub84B}

upd2^TOE.GS00171

viable | recessive

upd2^Δ3-62

Sterility

Allele

fertile

upd2^Δ

fertile | recessive

upd2^Δ3-62

Other Phenotypes

Allele

abnormal immune response | adult stage

upd2^Δ

abnormal starvation stress response | adult stage, with Scer\GAL4^ppl.PP

upd2^UAS.EGFP

decreased body size | adult stage

upd2^Δ3-62

decreased body size | adult stage, with Scer\GAL4^ppl.PP

upd2^NIG.5988R

decreased body size | larval stage, with Scer\GAL4^Cg.PA

upd2^NIG.5988R

decreased cell number | adult stage

upd2^Δ3-62

decreased cell size | adult stage

upd2^Δ3-62

decreased occurrence of cell division | adult stage

upd2^Δ

decreased size | adult stage | somatic clone, with Scer\GAL4^R42A07.-FRT

upd2^HMS00901

increased cell number | larval stage, with Scer\GAL4^Hand.Δ

increased cell number | larval stage, with Scer\GAL4^NP1029

upd2^HMS00901

short lived | nutrition conditional, with Scer\GAL4^Hml.Δ

upd2^UAS.EGFP

Phenotype manifest in

Allele

adult anterior midgut epithelium, with Scer\GAL4^NP0001

upd2^UAS.cUa

adult anterior midgut epithelium, with Scer\GAL4^NP5130, Scer\GAL80^ts.αTub84B

upd2^HMS00901

adult posterior midgut epithelium, with Scer\GAL4^NP0001

upd2^UAS.cUa

embryonic/larval crystal cell | increased number | larval stage, with Scer\GAL4^Hand.Δ

embryonic/larval crystal cell | increased number | larval stage, with Scer\GAL4^NP1029

upd2^HMS00901

embryonic/larval oenocyte | third instar larval stage

upd2^Δ3-62

intestinal stem cell, with Scer\GAL4^NP5130, Scer\GAL80^ts.αTub84B

upd2^HMS00901

wing

upd2^Δ3-62

wing, with Scer\GAL4^Bx-MS1096

upd2^UAS.EGFP

Orthologs

Downloads

Download All DIOPT Orthologs

Human Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Homo sapiens (Human) (0)

Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

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:upd2. Refer to their site for version information.

Paralogs

Downloads

Download All DIOPT Paralogs

Paralogs (via DIOPT v9.1)

Gene Symbol

Score

Source

Alignment

Drosophila melanogaster (Fruit fly) (1)

upd1

2 of 13

Human Disease Associations

FlyBase Human Disease Model Reports

leptin deficiency or dysfunction

Disease Ontology (DO) Annotations

Models Based on Experimental Evidence ( 2 )

Allele

Disease

Evidence

References

upd2^HMS00948

model of congenital leptin deficiency

CEA

(Beshel et al., 2017)

upd2^UAS.cUa

model of cancer

CEA with Ras85D^V12.UAS

(Wu et al., 2010)

Potential Models Based on Orthology ( 0 )

Human Ortholog

Disease

Evidence

References

Modifiers Based on Experimental Evidence ( 4 )

Allele

Disease

Interaction

References

upd2^HMS00901

ameliorates intestinal cancer

modeled by Sox21a⁶

(Zhai et al., 2015)

modeled by yki^{S111A.S168A.S250A.UAS.Tag:V5}

(Pranoto et al., 2023)

upd2^HMS00948

ameliorates intestinal cancer

modeled by yki^{S111A.S168A.S250A.UAS.Tag:V5}

(Pranoto et al., 2023)

upd2^Δ3-62

ameliorates cancer

modeled by Ras85D^V12.UAS, l(2)gl^unspecified

(Wu et al., 2010)

upd2^NIG.5988R

ameliorates cancer

modeled by Ras85D^V12.cUa.UAS, dlg1^RNAi.UAS.cUa

(Lodge et al., 2021)

ameliorates intestinal cancer

modeled by Sox21a⁶

(Zhai et al., 2015)

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

Interaction Browsers

View in FlyBase Interactions Browser View in MIST tool (external link)

Please see the Physical Interaction reports below for full details

RNA-RNA

Physical Interaction

Assay

References

upd2 - mir-bft

quantitative reverse transcription pcr

(Kim et al., 2017)

protein-protein

Physical Interaction

Assay

References

upd2 - Atg8a

pull down, autoradiography

(Madan et al., 2024)

upd2 - dome

anti tag coimmunoprecipitation, anti tag western blot

(Fisher et al., 2016)

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

upd2

suppressible

dome

(Hombria et al., 2005)

upd2

suppressible

Sox21a

(Zhai et al., 2015)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

Bub3

enhanceable

upd2|upd3

(Clemente-Ruiz et al., 2016)

Delta|twin

suppressible

upd2|upd3

(Vicente et al., 2018)

ebd1

suppressible

upd2

(Tian et al., 2017)

Edem1

suppressible

upd2

(Pathak and Varghese, 2021)

egr|CtBP

suppressible

upd2|upd3

(Worley et al., 2018)

egr|CtBP

suppressible

upd2

(Worley et al., 2018)

suppressible

upd2

(Zhai et al., 2015, Djiane et al., 2013)

N|yki

suppressible

upd2

(Pranoto et al., 2023)

PGRP-LE

suppressible

upd2|upd3

(Winkler et al., 2025)

PGRP-SD

suppressible

upd2

(Iatsenko et al., 2018)

suppressible

(Wu et al., 2010)

enhanceable

(Harris et al., 2020)

SH3PX1

suppressible

upd2|upd3

(Zhang et al., 2019)

Sox21a

suppressible

upd2

(Zhai et al., 2015)

upd1

suppressible

upd2

(Beshel et al., 2017)

enhanceable

(Wu et al., 2010)

enhanceable

(Osman et al., 2012)

yki

suppressible

upd2

(Pranoto et al., 2023)

External Data

Linkouts

DroID - A comprehensive database of gene and protein interactions.

FBgn0030904

MIST (genetic) - An integrated Molecular Interaction Database

32805

MIST (protein-protein) - An integrated Molecular Interaction Database

32805

Pathways

Signaling Pathways (FlyBase)

JAK-STAT SIGNALING PATHWAY CORE COMPONENTS

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.

Hippo signaling pathway - fly

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

Q9VWX2

Class of Gene

nuclear_gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

1-60

Cytogenetic map

17A3-17A3

Sequence location

X:18,240,654..18,243,284 [-]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

17A3-17A3

Limits computationally determined from genome sequence between P{EP}ari-1^EP317 and P{EP}EP1378

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

Experimentally Determined Recombination Data

Location

1-60

(FlyBase, 2016)

1-59.1

(Comeron et al., 2012)

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (12)

Aberrations including deletions of this gene Aberrations including duplications of this gene

Bloomington

55727

w^* upd2^Δ

55729

w^* upd2^Δ upd3^Δ

604936

w^* upd2^Δ; Df(4)CORL, PBac{WHr}CORL

67556

y¹ sc^* v¹ sev²¹; P{TOE.GS00171}attP40

33949

y¹ sc^* v¹ sev²¹; P{TRiP.HMS00901}attP2

33988

y¹ sc^* v¹ sev²¹; P{TRiP.HMS00948}attP2

81334

y¹ sc^* v¹ sev²¹; P{SAM.dCas9.GS03187}attP2

81323

y¹ sc^* v¹ sev²¹; P{SAM.dCas9.GS03176}attP40

NIG-Fly

5988R-1

P{NIG.5988R}1

5988R-3

P{NIG.5988R}3

HMS00901

y¹ sc^* v¹ sev²¹; P{TRiP.HMS00901}attP2

VDRC

v330691

P{VSH330691}attP40

Genomic Clones (16)

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.

FBgn0030904

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0030904

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Madan et al., 2024)

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

New stable cell line derived from S2-unspecified : S2-NP cells (expressing upd1, upd2 and upd3) and S2-NP-Stat92E cells were used.
(Kim et al., 2019)

Other Comments

upd2 and upd3 exhibit an additive effect in inducing intestinal stem cell proliferation in response to gut infection.

(Osman et al., 2012)

upd2 is a secreted factor produced by the fat body in response to dietary fat and sugars. upd2 activates JAK/STAT signaling in a population of GABAergic neurons, relieving their inhibitory effect on the insulin-producing cells and in turn resulting in the secretion of insulin-like peptides into the hemolymph to promote systemic growth and fat storage.

(Rajan and Perrimon, 2012)

Gene expression is increased in response to the presence of two copies of Scer\GAL4^hs.PB.

(Liu and Lehmann, 2008)

upd2 is not required for os-induced JAK/STAT signalling.

(Baeg et al., 2005)

S2-derived S2-NP cells transfected with dsRNA made from templates generated with primers directed against upd2 exhibit a reduction in JAK/STAT activity, indicating upd2 is involved in the positive regulation of JAK/STAT signalling in these cells.

(Baeg et al., 2005)

Relationship to Other Genes

Source for database merge of

Additional comments

"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: CG5988 upd2

(Agaisse and Perrimon, 2004)

Nomenclature comments

Etymology

Synonyms and Secondary IDs (18)

Reported As

Symbol Synonym

CG5988

(Meadows, 2018.10.31, Palu and Thummel, 2016, Wright et al., 2011, Ni et al., 2010.12.1, Baeg et al., 2007, Brown et al., 2006, Wawersik et al., 2005, Agaisse et al., 2003, Ringrose et al., 2003, Castelli-Gair Hombria and Brown, 2002, Hou et al., 2002)

UPD2

(He et al., 2025, Ismaila et al., 2024, Mahishi et al., 2024, Sanusi et al., 2024, Chowański et al., 2021, Toprak et al., 2020, Staats et al., 2018, Londraville et al., 2017, Ferrandon, 2013, Nässel et al., 2013, Arbouzova and Zeidler, 2006, Castelli-Gair Hombria, 2006)

Unpaireds

(Jasper, 2020)

Upd

(Zhai et al., 2023, Enomoto and Igaki, 2022, Casares and McGregor, 2021, Finger et al., 2021, Lam Wong and Verheyen, 2021, Surkova et al., 2021, Vidaurre and Chen, 2021, Horne-Badovinac, 2020, Morata and Calleja, 2020, Kim et al., 2019, Mehrotra and Deshpande, 2018, Mbodj et al., 2016)

Upd-2

(Alvarez-Ochoa et al., 2021)

Upd-like

(Wells et al., 2013)

Upd2

(Diaz et al., 2025, Hirooka et al., 2025, Krejčová and Bajgar, 2025, Li et al., 2025, Park et al., 2025, De Groef et al., 2024, Hersperger et al., 2024, Monticelli et al., 2024, Nigg et al., 2024, Sun et al., 2024, Wang et al., 2024, Khan et al., 2023, Kietz and Meinander, 2023, Kim et al., 2023, Williams et al., 2023, Benoit et al., 2022, Chopra et al., 2022, He et al., 2022, Neophytou and Pitsouli, 2022, Pratomo et al., 2022, Ratnaparkhi and Sudhakaran, 2022, Xu et al., 2022, Yu et al., 2022, Zhang and Edgar, 2022, Bilder et al., 2021, Chatterjee and Perrimon, 2021, De Groef et al., 2021, Dillard et al., 2021, García-López et al., 2021, Gillette et al., 2021, Harnish et al., 2021, Kim et al., 2021, Léopold, 2021, Lodge et al., 2021, Meschi and Delanoue, 2021, Montanari and Royet, 2021, Oliveira et al., 2021, Parra-Peralbo et al., 2021, Rosendo Machado et al., 2021, Schneider and Imler, 2021, Sciambra and Chtarbanova, 2021, Semaniuk et al., 2021, Yeom et al., 2021, Ahmad et al., 2020, Colombani and Andersen, 2020, Grenier and Leulier, 2020, Hao et al., 2020, Ingaramo et al., 2020, Kim et al., 2020, Koyama et al., 2020, Krautz et al., 2020, Lee et al., 2020, Lin and Hsu, 2020, Malita and Rewitz, 2020, Moore et al., 2020, Tafesh-Edwards and Eleftherianos, 2020, Texada et al., 2020, Zhou and Boutros, 2020, Zhou et al., 2020, Powers and Srivastava, 2019, Lehmann, 2018, Chng et al., 2017, Daisley et al., 2017, Kim et al., 2017, Liu and Jin, 2017, Mussabekova et al., 2017, Rajan et al., 2017, Zhao and Karpac, 2017, Zhou et al., 2017, Alfa and Kim, 2016, Bonfini et al., 2016, Droujinine and Perrimon, 2016, Fisher et al., 2016, Jiang et al., 2016, Katzenberger et al., 2016, Kubrak et al., 2016, Nässel and Vanden Broeck, 2016, Strigini and Leulier, 2016, Toggweiler et al., 2016, Vanha-Aho et al., 2016, Wakabayashi et al., 2016, Zheng et al., 2016, Figueroa-Clarevega and Bilder, 2015, Gold and Brückner, 2015, Pasco et al., 2015, Wang et al., 2015, Bausek and Zeidler, 2014, Guo et al., 2014, Huang et al., 2014, Owusu-Ansah and Perrimon, 2014, Wang et al., 2014, Andersen et al., 2013, Bausek, 2013, Hyun, 2013, Kannan and Fridell, 2013, Koyama et al., 2013, Mbodj et al., 2013, Morin-Poulard et al., 2013, Nässel et al., 2013, Rajan and Perrimon, 2013, Shim et al., 2013, Stec et al., 2013, Zeidler and Bausek, 2013, Zhou et al., 2013, Zoranovic et al., 2013, Amoyel and Bach, 2012, Kounatidis and Ligoxygakis, 2012, Liu et al., 2012, Apidianakis and Rahme, 2011, Biteau et al., 2011, Jiang et al., 2011, Stec and Zeidler, 2011, Wright et al., 2011, Zhao et al., 2011, Ren et al., 2010, Vidal et al., 2010, González et al., 2009, Jiang et al., 2009, Rivas et al., 2008, Pei and Harrison, 2007, Baeg et al., 2005, Zeidler et al., 2005)

Upd2/Leptin

(Handke et al., 2013)

cg5988

(Brown et al., 2003)

unpaired 2

(Wang et al., 2016)

upd

(Herrera and Bach, 2021, Jawkar and Nongthomba, 2020, Monahan and Starz-Gaiano, 2016, Martinez and Cavalli, 2010)

upd-2

(Jourjine, 2017, Roeder et al., 2009)

upd2

(Dong and Cheng, 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, Yuan et al., 2025, Zhou et al., 2025, Balakireva et al., 2024, Eslahi et al., 2024, Feng et al., 2024, Fioriti et al., 2024, Guo et al., 2024, Ju et al., 2024, Li et al., 2024, Li et al., 2024, Liu et al., 2024, Usman et al., 2024, Wang et al., 2024, Chen et al., 2023, Jiang et al., 2023, Li et al., 2023, Molina-Gil et al., 2023, Nagai et al., 2023, Neophytou et al., 2023, Petsakou et al., 2023, Tuo et al., 2023, Xu et al., 2023, Yamada et al., 2023, Yang et al., 2023, Zhou and Boutros, 2023, Cao et al., 2022, Evans et al., 2022, Gera et al., 2022, Keshav et al., 2022, Shen et al., 2022, Singh and Agrawal, 2022, Cai et al., 2021, Destalminil-Letourneau et al., 2021, Dong et al., 2021, Fabian et al., 2021, Gogia et al., 2021, Joy et al., 2021, Juarez-Carreño et al., 2021, Kim et al., 2021, Lourido et al., 2021, Maurya et al., 2021, Pathak and Varghese, 2021, Sanhueza et al., 2021, Ahmed et al., 2020, Chen et al., 2020, Cho et al., 2020, Dai et al., 2020, Delanoue and Romero, 2020, Geigges et al., 2020, Harris et al., 2020, Kierdorf et al., 2020, La Marca and Richardson, 2020, Maselko et al., 2020, Strilbytska et al., 2020, Strilbytska et al., 2020, Vizcaya-Molina et al., 2020, Wei et al., 2020, Wilson et al., 2020, Xu et al., 2020, Younes et al., 2020, Ahlers et al., 2019, Akmammedov et al., 2019, Asri et al., 2019, Bailetti et al., 2019, Banerjee et al., 2019, Gultekin and Steller, 2019, Herrera and Bach, 2019, Hill et al., 2019, Houtz et al., 2019, Lin et al., 2019, Lin et al., 2019, Saavedra and Perrimon, 2019, Si et al., 2019, Singh et al., 2019, Zhang et al., 2019, Ahmed-de-Prado et al., 2018, Bazzi et al., 2018, Gáliková and Klepsatel, 2018, Jia et al., 2018, Prange et al., 2018, Romey-Glüsing et al., 2018, Tokusumi et al., 2018, Vicente et al., 2018, Banerjee et al., 2017, Beshel et al., 2017, Kenmoku et al., 2017, Lee et al., 2017, Mattila and Hietakangas, 2017, Takemura and Nakato, 2017, Terriente-Félix et al., 2017, Tian et al., 2017, Transgenic RNAi Project members, 2017-, Xu et al., 2017, Zhang et al., 2017, Atkins et al., 2016, Guo et al., 2016, Hoi et al., 2016, Shen et al., 2016, Shih et al., 2016, Tian et al., 2016, Toggweiler et al., 2016, Wang et al., 2016, Ayyaz et al., 2015, Doggett et al., 2015, Gao et al., 2015, Katsuyama et al., 2015, Patel et al., 2015, Shapiro-Kulnane et al., 2015, Wagner et al., 2015, Woodcock et al., 2015, Yurgel et al., 2015, Zang et al., 2015, Zhai et al., 2015, Zhang et al., 2015, Kux and Pitsouli, 2014, Li et al., 2014, Djiane et al., 2013, Kannan and Fridell, 2013, Kemp et al., 2013, Zeidler and Bausek, 2013, Chakrabarti et al., 2012, Cordero et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Kelsey et al., 2012, Osman et al., 2012, Osman et al., 2012, Rajan and Perrimon, 2012, Feng et al., 2011, Karpac et al., 2011, Sinenko et al., 2011, Wright et al., 2011, Beebe et al., 2010, Shaw et al., 2010, Sotillos et al., 2010, Wu et al., 2010, Classen et al., 2009, Liu and Lehmann, 2008, López-Onieva et al., 2008, Pastor-Pareja et al., 2008, Wang and Harrison, 2008, Krzemień et al., 2007, Wang and Harrison, 2007, Hombria et al., 2005, Wawersik et al., 2005)

Name Synonyms

Unpaired 2

(De Groef et al., 2021, Chng et al., 2017, Mattila and Hietakangas, 2017)

Unpaired-2

(Jourjine, 2017, Castillo et al., 2015, Miyamoto et al., 2013)

Unpaired2

(Moraes and Montagne, 2021, Alfa and Kim, 2016, Sotillos et al., 2008)

unpaired 2

(Shafer, 2025, Yurgel et al., 2015, Owusu-Ansah and Perrimon, 2014, Piccirillo et al., 2014, Nässel et al., 2013, González et al., 2009, Hill-Burns and Clark, 2009, Hauenschild et al., 2008, Kula-Eversole and Hatini, 2007, Agaisse and Perrimon, 2004)

unpaired-2

(Kubrak et al., 2016)

upd homolog

(Feng et al., 2012)

upd2

(Tokusumi et al., 2018)

Secondary FlyBase IDs

Datasets (0)

Study focus (0)

Experimental Role

Project

Project Type

Title

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 28 )

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.

32805

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.

AAF48815

RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.

NM_001370039

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.

Q9VWX2

UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information

Q9VWX2

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.

Q9VWX2

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

32805/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0030904

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0030904

FlyMine - An integrated database for Drosophila genomics

FBgn0030904

InterPro - A database of protein families, domains and functional sites

Unpaired protein

KEGG Genes - Molecular building blocks of life in the genomic space.

dme:Dmel_CG5988

MARRVEL_MODEL - MARRVEL (model organism gene)

32805

Linkouts

Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones

FBgn0030904

DroID - A comprehensive database of gene and protein interactions.

FBgn0030904

DRSC - Results frm RNAi screens

FBgn0030904

Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.

upd2_1

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0030904

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0030904

Flygut - An atlas of the Drosophila adult midgut

FBgn0030904

Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution

Interactive Fly

KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.

Hippo signaling pathway - fly

MIST (genetic) - An integrated Molecular Interaction Database

32805

MIST (protein-protein) - An integrated Molecular Interaction Database

32805

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

Q9VWX2

References (431)

The following external sites may use different assemblies or annotations than FlyBase.

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