FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\Wnt4
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General Information
Symbol
Dmel\Wnt4
Species
D. melanogaster
Name
Wnt oncogene analog 4
Annotation Symbol
CG4698
Feature Type
protein_coding_gene
FlyBase ID
FBgn0010453
Gene Model Status
Current
Stock Availability
51 publicly available
Gene Summary
Binds as a ligand to a family of frizzled seven-transmembrane receptors and acts through a cascade of genes on the nucleus. Acts downstream of homeotic complex genes in the visceral mesoderm and is required for embryonic segmentation. Also required for cell movement and FAK regulation during ovarian morphogenesis. (UniProt, P40589)
Contribute a Gene Snapshot for this gene.
All Summaries
Also Known As

DWnt4, DWnt-4, Wnt-4

Key Links
Genomic Location
Cytogenetic map
27E7-27E8
Sequence location
Recombination map
2-25
RefSeq locus
NT_033779 REGION:7255419..7277171
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 (22 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
enables frizzled binding
(Wu and Nusse, 2002)
enables protein binding
inferred from physical interaction with FLYBASE:fj; FB:FBgn0000658
(Lim et al., 2005)
enables receptor ligand activity
inferred from genetic interaction with FLYBASE:otk; FB:FBgn0004839
(Tan et al., 2024)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
enables cytokine activity
inferred from biological aspect of ancestor with PANTHER:PTN000246517
(GO Reference Genome Project, 2011-)
enables frizzled binding
inferred from biological aspect of ancestor with PANTHER:PTN000246517
(GO Reference Genome Project, 2011-)
enables signaling receptor binding
inferred from electronic annotation with InterPro:IPR005817
(InterPro Project Members, 2004-)
inferred from sequence or structural similarity with FLYBASE:wg; FB:FBgn0284084
(Gene Ontology Curators, 2002-)
Biological Process (14 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
involved_in cell migration
inferred from mutant phenotype
(Cohen et al., 2002)
involved_in establishment of planar polarity
inferred from genetic interaction with FLYBASE:fj; FB:FBgn0000658
(Lim et al., 2005)
involved_in female gonad development
inferred from mutant phenotype
(Cohen et al., 2002)
involved_in germ-line stem-cell niche homeostasis
inferred from mutant phenotype
(Wang et al., 2015)
involved_in motor neuron axon guidance
inferred from mutant phenotype
(Inaki et al., 2007)
involved_in non-canonical Wnt signaling pathway
inferred from mutant phenotype
(Sato et al., 2006)
involved_in regulation of hemocyte proliferation
inferred from mutant phenotype
(Sinenko et al., 2010)
involved_in retinal ganglion cell axon guidance
inferred from mutant phenotype
(Sato et al., 2006)
involved_in salivary gland morphogenesis
inferred from mutant phenotype
(Harris and Beckendorf, 2007)
involved_in synaptic target inhibition
inferred from mutant phenotype
(Inaki et al., 2007)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
involved_in canonical Wnt signaling pathway
inferred from biological aspect of ancestor with PANTHER:PTN000246517
(GO Reference Genome Project, 2011-)
involved_in cell fate commitment
inferred from biological aspect of ancestor with PANTHER:PTN000246517
(GO Reference Genome Project, 2011-)
involved_in neuron differentiation
inferred from biological aspect of ancestor with PANTHER:PTN000246517
(GO Reference Genome Project, 2011-)
involved_in Wnt signaling pathway
inferred from electronic annotation with InterPro:IPR005817
(InterPro Project Members, 2004-)
inferred from sequence or structural similarity with FLYBASE:wg; FB:FBgn0284084
(Gene Ontology Curators, 2002-, Graba et al., 1995)
Cellular Component (3 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
located_in cell surface
inferred from direct assay
(Sato et al., 2006)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
located_in extracellular region
inferred from sequence model
(Graba et al., 1995)
non-traceable author statement
(Cohen et al., 2002)
inferred from sequence or structural similarity with FLYBASE:wg; FB:FBgn0284084
(Gene Ontology Curators, 2002-)
inferred from electronic annotation with InterPro:IPR005817
(InterPro Project Members, 2004-)
is_active_in extracellular space
inferred from biological aspect of ancestor with PANTHER:PTN000246517
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the Wnt family. (P40589)
Protein Signatures (InterPro)
Summaries
Gene Group (FlyBase)
WNTs -
WNTs are evolutionarily conserved secreted Cys-rich glycoproteins, defined by sequence homology to the original members of the family - Wnt1 in mouse and wingless (wg) in Drosophila. They are extracellular ligands for members of the Frizzled family of receptors as well as other receptors. (Adapted from PMID:23151663).
Protein Function (UniProtKB)
Binds as a ligand to a family of frizzled seven-transmembrane receptors and acts through a cascade of genes on the nucleus. Acts downstream of homeotic complex genes in the visceral mesoderm and is required for embryonic segmentation. Also required for cell movement and FAK regulation during ovarian morphogenesis.
(UniProt, P40589)
Summary (Interactive Fly)

ligand - Wnt homolog - mesodermal - a Wingless antagonist - transposon dysregulation modulates Wnt4 signaling to control germline stem cell differentiation

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

Please see the JBrowse view of Dmel\Wnt4 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 P40589)

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 5.47

Gene model reviewed during 5.55

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Wnt4-RA
FBtr0089291
NM_057624
3367
539
Wnt4-RB
FBtr0334799
NM_001273258
2938
539
Additional Transcript Data and Comments
Reported size (kB)

3.2 (northern blot)

(Graba et al., 1995)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
Wnt4-PA
FBpp0088345
58.7
539
9.29
P40589
NP_476972
AAF52499
Wnt4-PB
FBpp0306845
58.7
539
9.29
P40589
NP_001260187
AGB92723
Polypeptides with Identical Sequences

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

539 aa isoforms: Wnt4-PA, Wnt4-PB
Additional Polypeptide Data and Comments
Reported size (kDa)

389 (aa)

(Graba et al., 1995)
Comments
External Data
Post Translational Modification

Palmitoleoylated by porcupine. The lipid group functions as a sorting signal, targeting the ligand to polarized vesicles that transport Wnt4 to unique sites at the cell surface. Depalmitoleoylated by notum, leading to inhibit Wnt signaling pathway.

(UniProt, P40589)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
UniProt
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\Wnt4 using the Feature Mapper tool.

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

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

-0.32

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Graba et al., 1995)
presumptive embryonic/larval central nervous system
(Graba et al., 1995)
ectoderm
(Graba et al., 1995)
embryonic stage 7 -- 8
antennal anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
central brain anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
dorsal head epidermis anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
visual anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
foregut anlage
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
ventral nerve cord primordium

Comment: reported as ventral nerve cord anlage

(Fisher et al., 2012)
embryonic stage 8 -- 11
organism | segment polarity expression pattern
(Graba et al., 1995)
embryonic stage 9 -- 10
antennal primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
central brain primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
visual primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
dorsal head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
lateral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
ventral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
foregut primordium
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
ventral nerve cord primordium
(Fisher et al., 2012)
embryonic stage 10
cardiogenic mesoderm | dorsal | uniform
(Tauc et al., 2012)
embryonic stage 11
organism | segmentally repeated
(Sato et al., 1999)
parasegment 4 of visceral trunk mesoderm
(Sato et al., 1999)
larval ventral nerve cord
(Sato et al., 1999)
head segment | segmentally repeated
(Buratovich et al., 2000)
epidermis | segmentally repeated | ventral
(Buratovich et al., 2000)
visceral trunk mesoderm
(Chen et al., 2016)
embryonic stage 11 -- 12
central brain primordium
(Fisher et al., 2012)
clypeo-labral primordium
(Fisher et al., 2012)
dorsal epidermis primordium
(Fisher et al., 2012)
embryonic ganglion mother cell
(Fisher et al., 2012)
embryonic inner optic lobe primordium
(Fisher et al., 2012)
embryonic leading edge cell primordium

Comment: reported as embryonic leading edge cell specific anlage

(Fisher et al., 2012)
embryonic optic lobe primordium
(Fisher et al., 2012)
foregut primordium
(Fisher et al., 2012)
gnathal primordium
(Fisher et al., 2012)
hindgut proper primordium
(Fisher et al., 2012)
longitudinal trunk visceral muscle primordium
(Fisher et al., 2012)
procephalic neuroblast
(Fisher et al., 2012)
ventral epidermis primordium
(Fisher et al., 2012)
embryonic stage 11 -- 13
dorsal ectoderm | segmentally repeated
(Tauc et al., 2012)
embryonic stage 13
embryonic/larval heart
(Chen et al., 2016)
abdominal dorsal muscle cell
(Chen et al., 2016)
embryonic stage 13 -- 16
embryonic anal pad
(Fisher et al., 2012)
embryonic brain
(Fisher et al., 2012)
embryonic central brain neuron
(Fisher et al., 2012)
embryonic epipharynx
(Fisher et al., 2012)
embryonic esophagus
(Fisher et al., 2012)
embryonic foregut
(Fisher et al., 2012)
embryonic head epidermis
(Fisher et al., 2012)
embryonic head sensory system
(Fisher et al., 2012)
embryonic hindgut
(Fisher et al., 2012)
embryonic large intestine
(Fisher et al., 2012)
embryonic proventriculus
(Fisher et al., 2012)
embryonic rectum
(Fisher et al., 2012)
embryonic small intestine
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
larval nervous system
(Fisher et al., 2012)
gastric caecum
(Fisher et al., 2012)
lateral cord neuron
(Fisher et al., 2012)
larval ventral nerve cord
(Fisher et al., 2012)
embryonic optic lobe primordium
(Fisher et al., 2012)
embryonic inner optic lobe primordium
(Fisher et al., 2012)
embryonic heart cardioblast
(Chen et al., 2016)
embryonic heart ostial cardioblast
(Chen et al., 2016)
embryonic stage 14
embryonic/larval foregut
(Graba et al., 1995)
parasegment 4 of visceral trunk mesoderm
(Graba et al., 1995)
parasegment 8 of visceral trunk mesoderm
(Graba et al., 1995)
embryonic/larval hindgut
(Graba et al., 1995)
embryonic stage 15 -- 17
embryonic cardioblast
(Tauc et al., 2012)
embryonic stage 16
gastric caecum
(Graba et al., 1995)
embryonic midgut constriction 2
(Graba et al., 1995)
embryonic/larval heart
(Tauc et al., 2012)
muscle cell of ventral longitudinal muscle 1 | restricted
(Inaki et al., 2007)
muscle cell of A1-7 ventral longitudinal muscle 2 | restricted
(Inaki et al., 2007)
larval ostial pore
(Chen et al., 2016)
embryonic stage 16 -- 17
larval pericardial cell
(Tauc et al., 2012)
adult stage | female
germarium
(Mottier-Pavie et al., 2016)
Additional Descriptive Data

Wnt4 is expressed in visceral mesoderm at embryonic stage 11. At stage 13, after cardioblasts are specified, transcripts can be detected at a low level in cardioblasts and at high levels in certain posterior segment dorsal muscles. Wnt4 expression becomes stronger in the cardioblasts at stage 15, with higher expression levels in the posterior segments. Expression in the posterior three pairs of ostia progenitors increases dramatically just before ostia formation starts. Wnt4 expression increases progressively from anterior to posterior with the highest expression in the posterior-most pairs of ostia cells. At stage 16, Wnt4 expression is highest in the ostia cells and at low levels in non-ostia cardioblasts.

(Chen et al., 2016)

At embryonic stage 10, Wnt4 transcript is expressed in a continuous band along the dorsal side of the embryo, where heart progenitors are are specified. Wnt4 transcript is present in both the ectoderm and mesoderm at this stage. From embryonic stage 11, ectodermal expression becomes segmented, and is localized on the anterior side of en stripes; mesodermal expression is undetectable. Starting at stage 15, Wnt4 transcript is expressed myocardial cells, and stage 16 is more pronounced in the posterior region of the dorsal vessel. At the end of embryogenesis, Wnt4 transcript is expressed in mycardial and pericardial cells.

(Tauc et al., 2012)

Wnt4 transcripts are first detected in stage 8 embryos where they appear at the anterior tip and in a segment polarity pattern along the trunk. 15 stripes are observed in the posteriormost cells of each parasegment. They are one cell wide and overlap the ventralmost wg cells. Shortly after, expression is seen in the foregut primordium and in dorsal patches that are thought to correspond to cells of the labral and ocular segment primordia. By stage 14, transcripts are visible in the CNS and the developing gut. CNS expression is not uniform but increases, progressively from anterior to posterior regions of the embryo. In the gut, transcripts are seen in the foregut, hindgut, and in two regions of the visceral mesoderm corresponding to parasegments 4 and 8. The latter two sites correspond to the primordia of the gastric caeca and the second midgut constriction.

(Graba et al., 1995)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
pupal stage
ovarian sheath
(Cohen et al., 2002)
adult stage
germarium region 2
(Cohen et al., 2002)
germarium region 1
(Cohen et al., 2002)
ovarian sheath
(Cohen et al., 2002)
germarium cap cell
(Cohen et al., 2002)
Additional Descriptive Data

A 2 h after pupal formation (APF) protein is detected in the apical cells as they migrate basally. At 16 h APF Wnt4-expressing cells have migrated between the terminal filaments and developing germaria. At 18 h APF these cells have moved between individual basal stalk clusters. Expression is detected in sheath, basal half of the terminal filament cells and regions I and II of the germarium in the adult.

(Cohen et al., 2002)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cell surface
inferred from direct assay
(Sato et al., 2006)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Wnt4 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
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
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
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 7-8
  • Stages(s) 11-12
  • Stages(s) 13-16
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 ( 34 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Wnt4
Transgenic constructs containing regulatory region of Wnt4
Aberrations (Deficiencies and Duplications) ( 7 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal (with Df(2L)BSC324)
lethal | heat sensitive, with Scer\GAL4ptc-559.1
lethal - all die before end of P-stage, with Scer\GAL4ptc-559.1
partially lethal
partially lethal - majority die | recessive
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4tin.CΔ4
Sterility
Allele
sterile | recessive
Other Phenotypes
Allele
abnormal cell migration
abnormal cell shape
abnormal neuroanatomy
abnormal neuroanatomy, with Scer\GAL4NP6099
abnormal neuroanatomy (with Wnt4C1)
abnormal neuroanatomy (with Wnt4EMS23)
abnormal neuroanatomy (with Wnt4P23)
abnormal neuroanatomy | larval stage (with Wnt4C1)
abnormal neuroanatomy | larval stage (with Wnt4EMS23)
abnormal neuroanatomy | P-stage (with Wnt4EMS23)
abnormal neuroanatomy | P-stage (with Wnt4P23)
abnormal neuroanatomy | P-stage | somatic clone, with Scer\GAL4bsh.M.PT
abnormal pain response | adult stage, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
abnormal planar polarity, with Scer\GAL4dpp.blk1
abnormal planar polarity, with Scer\GAL4ptc-559.1
abnormal planar polarity (with Df(2L)spdj2)
abnormal planar polarity | somatic clone, with Scer\GAL4Act5C.PP
abnormal wound healing, with Scer\GAL4A58
some die during larval stage | recessive
some die during pharate adult stage, with Scer\GAL4ptc-559.1
visible, with Scer\GAL4dpp.blk1
visible, with Scer\GAL4ptc-559.1
visible | heat sensitive (with Df(2L)spdj2)
Phenotype manifest in
Allele
alar lobe | ectopic | heat sensitive, with Scer\GAL4ptc-559.1
anterior crossvein, with Scer\GAL4dpp.blk1
anterior crossvein, with Scer\GAL4ptc-559.1
axon | larval stage (with Wnt4C1)
axon | larval stage (with Wnt4EMS23)
dendrite, with Scer\GAL4A58
denticle | ectopic, with Scer\GAL4arm.PS
denticle | ectopic, with Scer\GAL4da.G32
dorsal denticle, with Scer\GAL4da.G32
dorsocentral bristle | increased number, with Scer\GAL4ptc-559.1
egg chamber
embryonic/first instar larval cuticle, with Scer\GAL4da.G32
embryonic/larval heart, with Scer\GAL4HCH.Hand
embryonic/larval heart (with Df(2L)BSC226)
embryonic/larval heart | embryonic stage 14
embryonic/larval optic stalk (with Wnt4C1)
embryonic/larval optic stalk (with Wnt4EMS23)
embryonic cardioblast, with Scer\GAL4HCH.Hand
embryonic cardioblast (with Df(2L)BSC226)
embryonic cardioblast | embryonic stage 14
embryonic epidermis | dorsal, with Scer\GAL4da.G32
embryonic somatic muscle cell
embryonic ventral epidermis, with Scer\GAL4arm.PS
eye equator | heat sensitive (with Df(2L)spdj2)
eye photoreceptor cell
eye photoreceptor cell, with Scer\GAL4NP6099
eye photoreceptor cell (with Df(2L)spdj2)
eye photoreceptor cell (with Wnt4C1)
eye photoreceptor cell (with Wnt4EMS23)
eye photoreceptor cell (with Wnt4P23)
eye ventral compartment (with Df(2L)spdj2)
germarium
haltere | ectopic | heat sensitive, with Scer\GAL4ptc-559.1
lamina
lamina, with Scer\GAL4NP6099
lamina (with Wnt4C1)
lamina (with Wnt4EMS23)
lamina (with Wnt4P23)
larval brain | larval stage (with Wnt4C1)
larval brain | larval stage (with Wnt4EMS23)
larval dorsal multidendritic neuron ddaE, with Scer\GAL4A58
larval dorsal multidendritic neuron ddaE | conditional, with Scer\GAL4A58
larval ostial pore
larval ostial pore, with Scer\GAL4HCH.Hand
larval ostial pore (with Df(2L)BSC226)
medulla | P-stage (with Wnt4EMS23)
medulla | P-stage (with Wnt4P23)
medulla | P-stage | somatic clone, with Scer\GAL4bsh.M.PT
medulla intrinsic neuron Mi1 | P-stage (with Wnt4EMS23)
medulla intrinsic neuron Mi1 | P-stage (with Wnt4P23)
medulla intrinsic neuron Mi1 | P-stage | somatic clone, with Scer\GAL4bsh.M.PT
mesonotum | heat sensitive, with Scer\GAL4ptc-559.1
microtubule, with Scer\GAL4A58
microtubule | conditional, with Scer\GAL4A58
neuron
neuron, with Scer\GAL4NP6099
neuron (with Wnt4C1)
neuron (with Wnt4EMS23)
neuron (with Wnt4P23)
ommatidium (with Df(2L)spdj2)
ovarian sheath
ovariole
ovary
photoreceptor cell R8 | P-stage (with Wnt4EMS23)
photoreceptor cell R8 | P-stage (with Wnt4P23)
posterior postalar bristle | increased number, with Scer\GAL4ptc-559.1
presumptive embryonic salivary gland
presumptive embryonic salivary gland, with Scer\GAL4bap.3
scutellar bristle | increased number, with Scer\GAL4ptc-559.1
scutellum | heat sensitive, with Scer\GAL4ptc-559.1
sternopleural bristle | increased number, with Scer\GAL4ptc-559.1
ventral denticle belt | ectopic, with Scer\GAL4da.G32
wing, with Scer\GAL4ptc-559.1
wing (with Df(2L)spdj2)
wing | ectopic | heat sensitive, with Scer\GAL4ptc-559.1
wing disc | heat sensitive, with Scer\GAL4ptc-559.1
wing hair, with Scer\GAL4dpp.blk1
wing hair, with Scer\GAL4en-e16E
wing hair, with Scer\GAL4ptc-559.1
wing hair | cell non-autonomous | somatic clone, with Scer\GAL4Act5C.PP
wing margin bristle | ectopic | heat sensitive, with Scer\GAL4ptc-559.1
wing sensillum | ectopic | heat sensitive, with Scer\GAL4ptc-559.1
wing vein | ectopic (with Df(2L)spdj2)
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) (19)
Hsap\WNT9B
8 of 14
Yes
Yes
1  
Hsap\WNT9A
6 of 14
No
Yes
1  
Hsap\WNT1
3 of 14
No
No
1  
Hsap\WNT6
3 of 14
No
No
Hsap\WNT11
3 of 14
No
Yes
Hsap\WNT16
3 of 14
No
No
Hsap\WNT2B
2 of 14
No
No
Hsap\WNT2
2 of 14
No
No
Hsap\WNT3A
2 of 14
No
No
Hsap\WNT3
2 of 14
No
No
Hsap\WNT4
2 of 14
No
No
Hsap\WNT5A
2 of 14
No
No
Hsap\WNT5B
2 of 14
No
No
Hsap\WNT7A
2 of 14
No
No
1  
Hsap\WNT7B
2 of 14
No
No
1  
Hsap\WNT8A
2 of 14
No
No
Hsap\WNT8B
2 of 14
No
No
Hsap\WNT10A
2 of 14
No
No
1  
Hsap\WNT10B
2 of 14
No
No
1  
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (19)
Rnor\Wnt9b
8 of 14
Yes
Yes
Rnor\Wnt9a
4 of 14
No
Yes
Rnor\Wnt6
3 of 14
No
No
Rnor\Wnt11
3 of 14
No
Yes
Rnor\Wnt16
3 of 14
No
No
Rnor\Wnt1
2 of 14
No
No
Rnor\Wnt2b
2 of 14
No
No
Rnor\Wnt3a
2 of 14
No
No
Rnor\Wnt3
2 of 14
No
No
Rnor\Wnt4
2 of 14
No
No
Rnor\Wnt5a
2 of 14
No
No
Rnor\Wnt5b
2 of 14
No
No
Rnor\Wnt7a
2 of 14
No
No
Rnor\Wnt7b
2 of 14
No
No
Rnor\Wnt2
1 of 14
No
No
Rnor\Wnt8a
1 of 14
No
No
Rnor\Wnt8b
1 of 14
No
No
Rnor\Wnt10a
1 of 14
No
No
Rnor\Wnt10b
1 of 14
No
No
Mus musculus (laboratory mouse) (19)
Mmus\Wnt9b
8 of 14
Yes
Yes
Mmus\Wnt9a
5 of 14
No
Yes
Mmus\Wnt1
3 of 14
No
No
Mmus\Wnt6
3 of 14
No
No
Mmus\Wnt11
3 of 14
No
Yes
Mmus\Wnt16
3 of 14
No
No
Mmus\Wnt2b
2 of 14
No
No
Mmus\Wnt2
2 of 14
No
No
Mmus\Wnt3a
2 of 14
No
No
Mmus\Wnt3
2 of 14
No
No
Mmus\Wnt4
2 of 14
No
No
Mmus\Wnt5a
2 of 14
No
No
Mmus\Wnt7a
2 of 14
No
No
Mmus\Wnt7b
2 of 14
No
No
Mmus\Wnt10a
2 of 14
No
No
Mmus\Wnt10b
2 of 14
No
No
Mmus\Wnt5b
1 of 14
No
No
Mmus\Wnt8a
1 of 14
No
No
Mmus\Wnt8b
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (20)
Xtro\wnt9a
3 of 13
Yes
Yes
Xtro\LOC493495
2 of 13
No
Yes
Xtro\wnt1
2 of 13
No
No
Xtro\wnt9b
2 of 13
No
Yes
Xtro\wnt11b
2 of 13
No
Yes
Xtro\wnt11
2 of 13
No
Yes
Xtro\ccdc78
1 of 13
No
Yes
Xtro\chmp4c
1 of 13
No
No
Xtro\gstk1
1 of 13
No
Yes
Xtro\LOC100487954
1 of 13
No
No
Xtro\mafa
1 of 13
No
No
Xtro\nek9
1 of 13
No
No
Xtro\nthl1
1 of 13
No
No
Xtro\wnt2
1 of 13
No
No
Xtro\wnt3a
1 of 13
No
No
Xtro\wnt3
1 of 13
No
No
Xtro\wnt5a
1 of 13
No
No
Xtro\wnt6
1 of 13
No
No
Xtro\wnt8a
1 of 13
No
No
Xtro\wnt10b
1 of 13
No
No
Danio rerio (Zebrafish) (24)
Drer\wnt9b
6 of 14
Yes
Yes
Drer\wnt9a
4 of 14
No
Yes
Drer\wnt6a
3 of 14
No
No
Drer\wnt6b
3 of 14
No
No
Drer\wnt11f2
3 of 14
No
Yes
Drer\wnt16
3 of 14
No
No
Drer\LOC568110
2 of 14
No
No
Drer\wnt1
2 of 14
No
No
Drer\wnt2ba
2 of 14
No
No
Drer\wnt2bb
2 of 14
No
No
Drer\wnt2
2 of 14
No
No
Drer\wnt3a
2 of 14
No
No
Drer\wnt3
2 of 14
No
No
Drer\wnt4
2 of 14
No
No
Drer\wnt7aa
2 of 14
No
No
Drer\wnt7ba
2 of 14
No
No
Drer\wnt7bb
2 of 14
No
No
Drer\wnt10a
2 of 14
No
No
Drer\wnt11
2 of 14
No
Yes
Drer\wnt4b
1 of 14
No
No
Drer\wnt5b
1 of 14
No
No
Drer\wnt8a
1 of 14
No
No
Drer\wnt8b
1 of 14
No
No
Drer\wnt10b
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (5)
Cele\mom-2
5 of 14
Yes
Yes
Cele\egl-20
3 of 14
No
No
Cele\cwn-2
2 of 14
No
No
Cele\lin-44
2 of 14
No
Yes
Cele\cwn-1
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (8)
Agam\AgaP_AGAP009735
10 of 12
Yes
Yes
Agam\AgaP_AGAP009731
3 of 12
No
No
Agam\AgaP_AGAP009733
3 of 12
No
No
Agam\AgaP_AGAP009734
3 of 12
No
No
Agam\AgaP_AGAP010283
3 of 12
No
No
Agam\AgaP_AGAP008678
2 of 12
No
No
Agam\AgaP_AGAP010290
2 of 12
No
No
Agam\AgaP_AGAP009732
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Wnt4. 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) (6)
wg
5 of 13
Wnt2
5 of 13
Wnt6
5 of 13
Wnt10
5 of 13
Wnt5
4 of 13
wntD
3 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 0 )
Allele
Disease
Evidence
References
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 3 )
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?
WNT9B; Wnt family member 9B
8 of 14
602864
       
      WNT9A; Wnt family member 9A
      6 of 14
      602863
           
          WNT1; Wnt family member 1
          3 of 14
          164820
           
          WNT6; Wnt family member 6
          3 of 14
          604663
              WNT11; Wnt family member 11
              3 of 14
              603699
                  WNT16; Wnt family member 16
                  3 of 14
                  606267
                      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
                      Wnt4|Wnt10|fz
                      enhanceable
                      fz2
                      (Han et al., 2020)
                      fz2|Wnt4|Wnt10
                      enhanceable
                      fz
                      (Han et al., 2020)
                      Wnt4
                      enhanceable
                      hep
                      (Sato et al., 2006)
                      Wnt4
                      enhanceable
                      Wnt5
                      (Harris and Beckendorf, 2007)
                      Starting gene(s)
                      Interaction type
                      Interacting gene(s)
                      Reference
                      dlp
                      suppressible
                      Wnt4
                      (Waghmare et al., 2020)
                      fj
                      enhanceable
                      Wnt4
                      (Lim et al., 2005)
                      fz
                      suppressible
                      Wnt4
                      (Lim et al., 2005)
                      hep
                      enhanceable
                      Wnt4
                      (Sato et al., 2006)
                      hh
                      suppressible
                      Wnt4
                      (Buratovich et al., 2000)
                      Nab2
                      suppressible
                      Wnt4
                      (Lee et al., 2020)
                      wg
                      suppressible
                      Wnt4
                      (Gieseler et al., 2001)
                      Wnt5
                      enhanceable
                      Wnt4
                      (Harris and Beckendorf, 2007)
                      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
                      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)
                      2L
                      Recombination map
                      2-25
                      Cytogenetic map
                      27E7-27E8
                      Sequence location
                      FlyBase Computed Cytological Location
                      Cytogenetic map
                      Evidence for location
                      27E7-27E8
                      Limits computationally determined from genome sequence between P{PZ}snRNP70K02107 and P{PZ}wg02657
                      Experimentally Determined Cytological Location
                      Cytogenetic map
                      Notes
                      References
                      27F-28A
                      (determined by in situ hybridisation)
                      (Gieseler et al., 1995)
                      Experimentally Determined Recombination Data
                      Location

                      2-25

                      (FlyBase, 2016)

                      2-23.9

                      (Comeron et al., 2012)
                      Left of (cM)
                      Right of (cM)
                      Notes

                      Maps close to wg.

                      (Graba et al., 1995)
                      Stocks and Reagents
                      Stocks (51)
                      Genomic Clones (29)
                      cDNA Clones (189)
                       

                      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

                      (Cohen et al., 2002)
                      Commercially Available Antibodies
                       
                      Cell Line Information
                      Publicly Available Cell Lines
                       
                        Other Stable Cell Lines
                         
                          Other Comments

                          Wnt4 is required for normal salivary gland migration in the embryo. It is required in the second phase of salivary gland migration, as the gland moves posteriorly within the embryo.

                          (Harris and Beckendorf, 2007)

                          Wnt4 regulates the dorsoventral specificity of retinal projections.

                          (Sato et al., 2006)

                          Wnt4 is required for cell movement and Fak56D regulation during ovarian morphogenesis. Wnt4 facilitates cell motility through regulation of focal adhesions.

                          (Cohen et al., 2002)

                          Wnt4 and wg produce distinct responses in cells of the dorsal embryonic epidermis. wg acts independently of hh signaling, while Wnt4 requires hh to elicit its effects.

                          (Buratovich et al., 2000)

                          Wnt4 has the capacity to antagonise wg signalling in the embryonic ventral epidermis.

                          (Gieseler et al., 1999)

                          wg and Wnt4 have partially overlapping embryonic expression patterns. Expression is regulated by HOM-C genes in the visceral mesoderm and segment polarity genes in the ectoderm. wg and Wnt4 achieve opposite, but complementary functions in intrasegmental cell patterning of the embryonic ectoderm.

                          (Gieseler et al., 1995)

                          Wnt4 has been characterised, Wnt4 gene product has a segment polarity-like expression pattern in the ectoderm and is controlled by homeotic complex (HOM-C) genes in the visceral mesoderm.

                          (Graba et al., 1995)
                          Relationship to Other Genes
                          Source for database merge of
                          Additional comments
                          Nomenclature History
                          Source for database identify of

                          Source for identity of: Wnt4 CG4698

                          (Deng, 1999.11)
                          Nomenclature comments
                          Etymology
                          Synonyms and Secondary IDs (16)
                          Reported As
                          Symbol Synonym
                          CG4698
                          (National Institute of Genetics Fly Stocks, 2022-, McEachern et al., 2014, Schertel et al., 2013, Keleman et al., 2009.8.5, Perkins et al., 2009)
                          DWnt-4
                          (Chen et al., 2008, Pandur et al., 2002, Cohen and Wilder, 2001, Cohen et al., 2001, Logan et al., 2001, Buratovich et al., 2000, Cohen et al., 2000, Wilder and Wallace, 2000, Wilder and Wallace, 2000, Buratovich and Wilder, 1999, Gieseler et al., 1999, Wilder et al., 1999, Buratovich and Wilder, 1998, Perrimon and Nusse, 1998, Cadigan and Nusse, 1997, Graba et al., 1997, Wilder et al., 1997, Wilder et al., 1997, Graba et al., 1995)
                          DWnt4
                          (Han et al., 2020, Yu et al., 2020, Port et al., 2008, Coudreuse and Korswagen, 2007, Sato et al., 2006, Lim et al., 2005, Sato et al., 2005, Gupta and Schupbach, 2003, Cadigan, 2002, Cohen and Wilder, 2002, Cohen et al., 2002, Peifer and McEwen, 2002, Wu and Nusse, 2002, Gieseler et al., 2001, Llimargas and Lawrence, 2001, Cox et al., 2000, Rizkalla and Siegfried, 2000, Reifegerste and Moses, 1999, Sato et al., 1999, Mariol et al., 1997, Gieseler et al., 1995, Gieseler et al., 1995, Mariol et al., 1995)
                          Dm DWnt4
                          (Konikoff et al., 2010)
                          Dwnt-4
                          (Swarup and Verheyen, 2012)
                          Dwnt4
                          (Perochon et al., 2018, Joshi et al., 2007, Bovolenta et al., 2006, Terry et al., 2006, Huang and Klein, 2004, Cadigan et al., 2002, Taylor, 2000)
                          WNT4
                          (Riahi et al., 2021, Harris et al., 2016)
                          Wnt
                          (Blair, 1999)
                          Wnt-4
                          (Tan et al., 2024, Bajpai et al., 2004, Jockusch and Ober, 2000, Wehrli and Tomlinson, 1998)
                          Wnt4
                          (Janssens et al., 2025, Cui et al., 2024, Goins et al., 2024, Liang and Luan, 2024, Corthals et al., 2023, Huang et al., 2023, Jetti et al., 2023, Deshpande et al., 2022, Kotian et al., 2022, Kubrak et al., 2022, Mieszczanek et al., 2022, Tarikere et al., 2022, Velten et al., 2022, Destalminil-Letourneau et al., 2021, Everetts et al., 2021, Feng et al., 2021, Joshi et al., 2021, Joy et al., 2021, Malin and Desplan, 2021, McLaughlin et al., 2021, Michki et al., 2021, Shin et al., 2021, Slaidina et al., 2021, Watanabe and Riddle, 2021, Cui et al., 2020, Rushton et al., 2020, Rust et al., 2020, Slaidina et al., 2020, Tu et al., 2020, Waghmare et al., 2020, Zhang and Cai, 2020, Hall et al., 2019, Story et al., 2019, Wu et al., 2019, Cho et al., 2018, Jankovics et al., 2018, Ripp et al., 2018, Tian et al., 2018, Waghmare and Page-McCaw, 2018, Hessinger et al., 2017, Suresh et al., 2017, Transgenic RNAi Project members, 2017-, Chen et al., 2016, Dominguez et al., 2016, Gene Disruption Project members, 2016-, Mottier-Pavie et al., 2016, Pearson et al., 2016, Wang et al., 2015, Hamada-Kawaguchi et al., 2014, Kerr et al., 2014, Linnemannstöns et al., 2014, Toshima et al., 2014, Schertel et al., 2013, Sopko and Perrimon, 2013, Webber et al., 2013, Wu et al., 2013, Herr and Basler, 2012, Japanese National Institute of Genetics, 2012.5.21, Kuroda et al., 2012, Lahaye et al., 2012, Nose, 2012, Cherbas et al., 2011, Peradziryi et al., 2011, Biehs et al., 2010, Bina et al., 2010, Inaki et al., 2010, Kong et al., 2010, Chen et al., 2008, Christensen and Cook, 2007.5.8, Christensen et al., 2007.10.29, Harris and Beckendorf, 2007, Inaki et al., 2007, Zeitlinger et al., 2007, Zeitouni et al., 2007, Casal et al., 2006, Christensen and Cook, 2006.12.5, Ma et al., 2006, Pearson et al., 2005)
                          anon-EST:Liang-2.4
                          clone 2.4
                          (Liang and Biggin, 1998)
                          dWnt4
                          (Bischoff et al., 2025, Humphries and Mlodzik, 2018)
                          wnt-4
                          (Liang and Biggin, 1998)
                          wnt4
                          (El-Danaf et al., 2025, Chen et al., 2022, Nguyen et al., 2022, Ewen-Campen et al., 2020, Mele and Johnson, 2019, Ewen-Campen et al., 2017, Luo et al., 2015, Khan et al., 2013, Sinenko et al., 2010, Srahna et al., 2006, Veeman et al., 2003, Bhat, 1999, Bhat, 1998)
                          Name Synonyms
                          Wnt oncogene analog 4
                          (McEachern et al., 2014)
                          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 ( 47 )
                            Sequence Crossreferences
                            NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
                            GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
                            GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
                            RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
                            UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
                            UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
                            UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
                            Other crossreferences
                            AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
                            BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
                            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
                            FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
                            iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
                            Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                            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 (229)