FB2025_05 , released December 11, 2025
Gene: Dmel\wg
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General Information
Symbol
Dmel\wg
Species
D. melanogaster
Name
wingless
Annotation Symbol
CG4889
Feature Type
FlyBase ID
FBgn0284084
Gene Model Status
Stock Availability
Gene Summary
wingless (wg) is a segment polarity gene that encodes a ligand of the Wnt/Wg signaling pathway. Its post-translational modification (addition of palmitoleate by the product of por) is essential for signaling activity. It contributes to segment polarity, tissue growth and patterning, neuromuscular junction morphogenesis, gut homeostasis and long term memory formation. [Date last reviewed: 2019-03-21] (FlyBase Gene Snapshot)
Also Known As

Wnt, Sp, Wnt-1, Gla, Sternopleural

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
2-26
RefSeq locus
NT_033779 REGION:7307159..7316265
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (80 terms)
Molecular Function (9 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from physical interaction with FLYBASE:fz2; FB:FBgn0016797
inferred from physical interaction with UniProtKB:Q9VVX3
inferred from direct assay
inferred from direct assay
inferred from physical interaction with UniProtKB:Q95ST2
inferred from physical interaction with UniProtKB:Q9VWV9
inferred from physical interaction with UniProtKB:Q95ST2
inferred from physical interaction with UniProtKB:Q8N474
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000246517
inferred from biological aspect of ancestor with PANTHER:PTN000246517
inferred from electronic annotation with InterPro:IPR005817
Biological Process (61 terms)
Terms Based on Experimental Evidence (58 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
involved_in dorsal closure
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
involved_in heart development
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:fz2; FB:FBgn0016797
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
involved_in long-term memory
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:Sox15; FB:FBgn0005613
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:Sox15; FB:FBgn0005613
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000246517
inferred from biological aspect of ancestor with PANTHER:PTN000246517
inferred from biological aspect of ancestor with PANTHER:PTN000246517
inferred from electronic annotation with InterPro:IPR005817
Cellular Component (10 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
located_in cell surface
inferred from direct assay
colocalizes_with early endosome
inferred from direct assay
inferred from direct assay
colocalizes_with late endosome
inferred from direct assay
located_in membrane raft
inferred from direct assay
inferred from direct assay
located_in plasma membrane
inferred from direct assay
located_in terminal bouton
inferred from direct assay
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR005817
is_active_in extracellular space
inferred from biological aspect of ancestor with PANTHER:PTN000246517
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the Wnt family. (P09615)
Summaries
Gene Snapshot
wingless (wg) is a segment polarity gene that encodes a ligand of the Wnt/Wg signaling pathway. Its post-translational modification (addition of palmitoleate by the product of por) is essential for signaling activity. It contributes to segment polarity, tissue growth and patterning, neuromuscular junction morphogenesis, gut homeostasis and long term memory formation. [Date last reviewed: 2019-03-21]
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).
Pathway (FlyBase)
WNT-TCF SIGNALING PATHWAY CORE COMPONENTS -
The canonical Wnt signaling pathway is initiated by the binding of a Wnt ligand to a frizzled family receptor on the cell surface. Activation of the pathway leads to the inhibition of cytoplasmic β-catenin (arm) degradation and its subsequent accumulation in the nucleus, where it regulates the transcription of target genes. (Adapted from FBrf0218499 and FBrf0223299).
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. Segment polarity protein. May be a growth factor. Acts on neighboring cells to regulate at least one gene, the homeobox segmentation gene engrailed. Wg signal represses arm phosphorylation. Wg signaling operates by inactivating the sgg repression of engrailed autoactivation. Wg and Wnt2 have a role in the developing trachea and together are responsible for all dorsal trunk formation. Wg also acts in the developing epidermis. Acts as a morphogen, and diffuses long distances despite its lipidation. Lipophorin is required for diffusion, probably by acting as vehicle for its movement, explaining how it can spread over long distances despite its lipidation. In non-neuronal cells, wls directs wg secretion via clathrin-mediated endocytosis and the retromer complex (a conserved protein complex consisting of Vps26 and Vps35) to sustain a wls traffic loop encompassing the Golgi, the cell surface, an endocytic compartment and a retrograde route leading back to the Golgi. In neuronal cells (the larval motorneuron NMJ), wg signal moves across the synapse through the release of wls-containing exosome-like vesicles.
(UniProt, P09615)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
Gla: Glazed
Eye reduced to one-fourth normal area and narrowed to a point ventrally. Eye color generally diluted but with some black patches. Ommatidia coalesce into gleaming, smooth sheet. Malpighian tubes of larva somewhat lighter than wild type; difficult to classify (Brehme and Demerec, 1942, Growth 6: 351-56). Abdomen of heterozygous female fails to distend with eggs; fertility impaired (Craymer, 1980, DIS 55: 200). Homozygous lethal. RK2A.
Sp: Sternopleural
thumb
Sp: Sternopleural
Edith M. Wallace, unpublished.
Sternopleural bristles increased in number. At 19, wild type; at 25, overlaps wild type; at 28-30, no overlap. Apparently does not affect sternopleural bristles on metathoracic segment converted by bx to a mesothoracic segment (Waddington, 1939, Growth Suppl. 1, pp. 37-44). Homozygous lethal. RK2.
spd: spade
Wings short and broad, pointed at tip, and warped at base. Effect on wing shape arises from excessive contraction of epithelium from inflated stage onward (Waddington, 1940, J. Genet. 41: 75-139). Overlaps wild type in existing stock. RK3.
spdfg: spade-flag
Wings about two-thirds the length and three-fourths the width of wild type, held tentlike over abdomen. Alulae absent or vestigial; proximal posterior wing margins often irregular with tendency to fold under about vein L4. Venation usually normal with occasional blistering. spdfg/spd has phenotype varying from slight shortening of wings to a shape midway between the two homozygotes. Excellent viability and fertility. RK1.
wg: wingless
The wg gene is involved both in controlling the segmentation pattern of embryos by affecting the posteriormost cells of each parasegment (Baker, 1987) and in controlling the imaginal disk pattern of the meso- and meta-thoracic segments that develop into wing, halter, and notum in pupae and adults (Sharma, 1973; Sharma and Copra, 1976; Morata and Lawrence, 1977). The gene is believed to control segment organization through an intercellular signaling mechanism (Baker, 1987, 1988b; Cabrera, Alonso, Johnston, Phillips, and Lawrence, 1987, Cell 50: 659-63; Ryjsewijk et al., 1987; Martinez-Arias et al., 1988). Mutants may be viable as adults or lethal as embryos or pupae. In embryonic lethal alleles (Babu, 1977; Nusslein-Volhard and Wieschaus, 1980), each segment shows a mirror-image duplication of the denticle bands at the expense of naked cuticle so that a continuous sheet of denticles (instead of repeated denticle bands) is produced (Cabrera et al., 1987). Dorsal abnormalities are more extreme than ventral ones, the dorsal cuticle being greatly reduced and covered by fine hairs (Baker, 1988a). These embryos lack head structures and filzkorper (Perrimon and Mahowald, 1987, Dev. Biol. 119: 587-600). In the nervous system, a single neuron, RP2, is missing; other neurons in the lineage are normal (Patel, Schafer, Goodman, and Holmgren, 1989, Genes Dev. 3: 890-904). The temperature-sensitive period for wgl-12, a heat-sensitive allele that is lethal at 25 (Baker, 1988a; Mohler, 1988), lies between gastrulation and the beginning of dorsal closure (11 hours after egg laying at 25). In pupal lethal and adult viable alleles, the ready-to-emerge pupae and the adults lack one or both wings and/or halteres, and there is a corresponding duplication of the meso- and metanota (Sharma, 1973; Sharma and Copra, 1976; Morata and Lawrence, 1977; Deak, 1978). This adult phenotype shows incomplete penetrance and variable expressivity and is affected by the ability of the wingless gene to function during the larval period (Baker, 1988a). Low temperature fails to rescue heteroallelic combinations of wg1 or wgl-18 with the heat-sensitive allele wgl-12 after the larval stages. Lethal as well as viable wg alleles are not cell-autonomous in adult mosaics (Babu and Bhat, 1986; Morata and Lawrence, 1977; Baker, 1988a).
Summary (Interactive Fly)

ligand - wnt family - segment polarity gene - plays a primary role in specifying the wing primordium, and a subsequent role mediating the patterning activities of the dorso-ventral compartment boundary - post-translational modification (addition of palmitoleate by Porcupine) is essential for signaling activity - contributes tissue growth and patterning, neuromuscular junction morphogenesis, gut homeostasis and long term memory formation.

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

Please see the JBrowse view of Dmel\wg 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 P09615)

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 5.44

Gene model reviewed during 5.51

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0079432
2919
468
Additional Transcript Data and Comments
Reported size (kB)

3.2 (northern blot)

3.0 (compiled cDNA)

3.0 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0079060
52.0
468
9.19
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)

468 (aa); 52.766 (kD predicted)

Comments
External Data
Subunit Structure (UniProtKB)

Monomer; folds by intramolecular disulfide bonds (PubMed:11821428). Interacts with porcupine (por) (PubMed:11821428). Interacts with wls; in the Golgi (PubMed:18193037). Interacts with en (PubMed:1335365). Interacts with the proteoglycan Cow (heparan sulfate-bound form); this stabilizes wg and promotes its extracellular distribution (PubMed:25360738). Interacts with peg; the interaction facilitates short-range diffusion of wg (PubMed:34580289).

(UniProt, P09615)
Post Translational Modification

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

Major form is glycosylated at 2 sites, glycosylation is stimulated by porcupine at the ER.

(UniProt, P09615)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\wg 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).

-0.28

Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
dorsal ectoderm anlage

Comment: anlage in statu nascendi

ectoderm anlage

Comment: anlage in statu nascendi

mesectoderm anlage

Comment: anlage in statu nascendi

mesoderm anlage

Comment: anlage in statu nascendi

ventral ectoderm anlage

Comment: anlage in statu nascendi

visual anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

antennal anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

dorsal head epidermis anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

antennal primordium

Comment: reported as procephalic ectoderm primordium

central brain primordium

Comment: reported as procephalic ectoderm primordium

visual primordium

Comment: reported as procephalic ectoderm primordium

dorsal head epidermis primordium

Comment: reported as procephalic ectoderm primordium

lateral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

ventral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

wg transcript distribution follows a similar pattern to the protein. Transcripts are first detected at early nuclear cycle 14 and remain strong through germband extension. They are first detected in the posterior region, starting slightly before anterior expression and posterior wg protein expression. At mid nuclear cycle 14, levels of the posterior band have increased and transcripts for segmental bands begin to appear in anterior segments. In early gastrulation, each segment has a thin stripe of wg mRNA and the posterior wg band is most intense. At stage 7, the posterior stripe remains strong and moves dorsally to form the "midgut plate". At stage 10, the posterior tissue expressing wg invaginates to later form the hindgut.

wg transcript is detected in the late third instar larval wing disc in a stripe that corresponds to the future wing margin and in two concentric rings that encircle the wing pouch. The inner wing gives rise to the distal hinge structures and the outer ring gives rise to proximal hinge structures. An additional stripe of wg expression marks the future mesonotum.

At 5 h after egg laying (AEL), wg is expressed in two rings of the hindgut, the inner covering the presumptive tubule primordia.

In early embryos wg transcript is expressed in two anterior domains and one broad posterior domain.

Expression of wg is seen in several domains of the wing disc, including two rings surrounding the wing pouch, the inner of which develops into the more distal stripe running through the adult wing hinge.

en and wg are expressed in imaginal discs from the time the imaginal disc cells segregate from the larval epidermis until the end of the third larval instar. wg is generally expressed in the anterior portions of the imaginal discs.

wg transcription is repressed by ectopic eve protein in evehs.PS embryos. Timing suggests that eve is a direct regulator of wg.

wg expression is seen on the ventral side of the proctodeum and then in the primordia of the Malphigian tubules as they evert from the proctodeum. As the tubules grow, wg is associated with the posterior side of each tubule.

wg is expressed in parasegment 8 in the mesoderm. This expression is abolished in abd-A, Ubx, and dpps4 homozygotes.

wg transcripts are most abundant in 3-6hr embryos and in pupae and are detectable at all other stages tested. wg transcripts are first detected in blastoderm embryos at the anterior pole and in a ring around the posterior end. They accumulate in a series of stripes, one per metameric unit in the extended germband. At germ band shortening, the stripes are 3-5 cells wide and include the most posterior cells of the anterior compartment of each segment. wg transcripts are also detected in the CNS, hindgut, procephalic lobe, labrum, and the analia.

The wg transcript is expressed in 16 evenly spaced stripes 2-3 cells wide in germ band extended embryos. Additional hybridization is detected in the procephalic lobe and the anterior head region.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

At the larval neuromuscular junction, wg protein is localized at large type Ib synaptic boutons in a dynamic pattern of punctuate distribution at the synaptic interface between motor neuron and muscle.

wg protein is detected in the intestinal stem cells, enteroblasts, and enteroendocrine cells in the posterior midgut epithelium.

The first expression of wg protein appears during cellularization of the blastoderm. At mid nuclear cycle 14, posterior wg appears in a band at 10% egg length. Two patches of anterior expression at 85% and 100% egg length are seen. This is before the appearance of any segmental stripes. As blastoderm stage progresses, the posterior stripe becomes stronger and remains the most prominent region of wg expression during gastrulation and early germ band extension. The segmental stripes appear sequentially from anterior to posterior during late cell cycle 14 blastoderm stage. During early gastrulation (stage 6), all 14 of the segmental stripes are formed and a strong posterior band of wg remains at 10% egg length. By late gastrulation, the posterior wg band migrates to a region called the midgut plate, which invaginates at the extended germ band and eventually forms the hindgut. wg is found in the future hindgut. The expression of the posterior band of wg was compared to other gap and pair-rule genes. wg is expressed subsequent to hb protein and appears during cellularization of the blastoderm in a narrow band posterior to the posterior hb stripe. Kr is first detected during late syncytial blastoderm but wg posterior expression just posterior to the Kr domain appears at mid nuclear cycle 14. eve antigen is first detected at the early blastoderm stage. wg posterior band expression occurs at mid nuclear cycle 14 posterior to the seventh eve stripe.

wg is expressed in all dorsal and ventral abdominal histoblast nest in segments A1-7 in females at 26hr APF. In males, expression is absent in segment A7 at 26hr APF.

At 0 and 5 hours after the second to third instar larval transition, wg is expressed in a stripe in the presumptive wing margin. At 10 hours, a ring of wg appears around the wing pouch. A faint patch of wg is visible in the notum at 15 hours and is more clearly resolved by 25 hours.

wg protein is not detected in embryonic lymph glands; expression is first detected in majority of lymph gland cells in newly-hatched first instar larvae, and is uniformly expressed in hematopoietic cells of the lymph gland through mid-second instar. At mid-to-late second instar, wg protein expression is down-regulated in the forming cortical zone of the lymph gland; a drop in wg expression in differentiating hemocytes occurs prior to those cells expressing the hemocyte marker Hml. wg protein expression is maintained in prohemocytes in the medullary zone through late third larval instar; expression is also observed in crystal cells in the cortical zone of the lymph gland. Late in the third larval instar, a second wave of wg protein expression occurs in mature Hml-expressing hemocytes.

wg protein accumulates between the luminal surface of the circular muscles and the basement membrane of the gut epithelia. Weak accumulation is also seen in intestinal stem cells.

In third instar wing disc, wg is expressed distally

in a stripe of cells that will form the adult wing margin and, proximally, in the inner and outer rings. The limit of wg expression in the inner ring coincides proximally with the proximal limit of rn expression and distally with both the distal limit of zfh2 and the proximal limits of dve and nab expression.

wg is expressed in the anterior and posterior boundary cells of the embryonic proventriculus.

At embryonic stage 16, wg is expressed in svp-expressing cells in segments A5-A7. At 30-36 hr APF, wg is expressed transiently in svp-expressing cells in segment A1-A5.

wg protein isdetected in two concentric rings toward the edges of the wing disc and in a stripe along the dorsal/ventral boundary of the wing pouch.

wg protein is detected in the procephalic neurectoderm from stage 8 onward in a domain spanning a broad area of the ocular and anterior antennal segment. Additional domains of wg expression include a small spot of expression in intercalary segment and a expression in the dorsal hemispheres of the clypeo-labral segment. 25% of the neuroblasts in the protocerebrum are wg positive as well as 3 neuroblasts in the deutocerebrum and a single neuroblast in the tritocerebrum.

Expression in procephalic neuroblasts stage 9-11: tritocerebrum - d4; deuterocerebrum - d1, d7, d8; protocerebrum - cd1, cd3, cd6, cd7, cd10-13, pd1, pd3, pd4, pd6, pd7, pd9, pd12, pd13

Strong wg protein expression was observed in glutamatergic type 1b synaptic boutons at the larval neuromuscular junction. The protein was observed both pre- and post-synaptically and evidence indicates that it is secreted from the pre-synaptic neuron and taken up by the post-synaptic muscle cell.

Using conventional staining techniques wg protein is detected in a stripe on the apical side of the presumptive notum. However, when an inactive form of fz2 that is still able to bind the wg protein is overexpressed in the underlying mesoderm, wg protein can be detected in the mesodermal tissue indicating that wg can diffuse across germ layers.

The wg protein is expressed in a specified subset of neuroblasts in embryonic stages 8-11. (see also FBrf 49374)

The expression pattern of wg protein in wghs.P embryos, 3 hours after heat shock, is similar to that in nkd mutant embryos.

wg expression was observed in 5 regions which are anterior to the centers of en expression. These are the "wg antennal stripe", the "wg head blob", the "wg intercalary spot", the "wg expression in the foregut" and the "wg labral spot". The relative positioning of the wg- and en-expressing cells was followed.

wg protein expression is seen on the ventral side of the proctodeum and then in the primordia of the Malpighian tubules as they evert from the proctodeum. As the tubules grow, wg protein is associated with the posterior side of each tubule.

The wg protein is expressed in the embryo in a each parasegment, in a 4-5 cell width stripe, just anterior to en expressing cells. Electron microscopy revealed that the wg protein accumulates in the cytoplasm of the wg expressing cells, and is then detected in the ECM and in en expressing cells. It seems that the wg protein is passed directly from cell to cell. wg protein is also detected up to two cell widths away from wg expressing cells.

In strong arm alleles, wg RNA is no longer detectable in embryos by late stage 9.

wg protein does not accumulate in the visceral mesoderm in embryos lacking abd-A. Its expression pattern is unchanged in Ubx mutants and in embryos with ectopic Ubx expression. wg expression is severely reduced in embryos lacking Ubx and its surrounding regulatory regions.

Marker for
Subcellular Localization
CV Term
Evidence
References
located_in cell surface
inferred from direct assay
colocalizes_with early endosome
inferred from direct assay
inferred from direct assay
colocalizes_with late endosome
inferred from direct assay
located_in membrane raft
inferred from direct assay
inferred from direct assay
located_in plasma membrane
inferred from direct assay
located_in terminal bouton
inferred from direct assay
Expression Deduced from Reporters
Reporter: P{en1}wg17en40cP1
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{en1}wgen11
Stage
Tissue/Position (including subcellular localization)
Reference
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GAL4-wg.M}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GalW}wgND382
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GawB}MD758
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GawB}wgGal4
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{PZ}wgSp-revP
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{wg(1.2)lacZ}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{wg-lacZ.nls}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{WLZ2.5L}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{WLZ4.5L}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{WLZΔG}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: TI{GAL4}wgKO.GAL4
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\wg in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
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
Images
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 141 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 165 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of wg
Transgenic constructs containing regulatory region of wg
Aberrations (Deficiencies and Duplications) ( 28 )
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
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
bouton & microtubule
dorsal mesothoracic disc & peripodial epithelium | somatic clone, with Scer\GAL4αTub84B.PL
dorsal mesothoracic disc & peripodial epithelium | somatic clone | cell non-autonomous, with Scer\GAL4αTub84B.PL
embryonic/larval dorsal vessel & embryonic myoblast
embryonic/larval somatic muscle & embryonic myoblast | dorsal
embryonic leading edge cell & actin filament
embryonic leading edge cell & actin filament, with Scer\GAL4ptc-559.1
embryonic leading edge cell & filopodium
embryonic leading edge cell & microtubule
embryonic leading edge cell primordium & microtubule
larval hindgut & ectoderm
leg & cuticle | dorsal
macrochaeta & scutum
Malpighian tubule & cell | conditional ts
mesothoracic pleurum & microchaeta
microchaeta & scutum
microchaeta & tarsal segment 5, with Scer\GAL4Dll-em212
mitochondrion & bouton
sensory mother cell & dorsal mesothoracic disc | ectopic, with Scer\GAL4sd-SG29.1
wing & epidermis
wing & macrochaeta
wing & macrochaeta, with Scer\GAL4Bx-MS1096
wing & macrochaeta, with Scer\GAL4C-765
wing & macrochaeta, with Scer\GAL4sd-SG29.1
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (19)
12 of 14
Yes
Yes
1  
5 of 14
No
No
4 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1  
2 of 14
No
No
1  
2 of 14
No
No
2 of 14
No
No
1  
2 of 14
No
No
1  
2 of 14
No
No
1  
2 of 14
No
No
1  
2 of 14
No
No
2 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (19)
13 of 14
Yes
Yes
3 of 14
No
No
3 of 14
No
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Mus musculus (laboratory mouse) (19)
12 of 14
Yes
Yes
5 of 14
No
No
4 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (23)
11 of 13
Yes
Yes
3 of 13
No
No
2 of 13
No
Yes
2 of 13
No
No
2 of 13
No
No
2 of 13
No
Yes
2 of 13
No
Yes
2 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Danio rerio (Zebrafish) (25)
13 of 14
Yes
Yes
3 of 14
No
No
3 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (5)
5 of 14
Yes
Yes
3 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (8)
12 of 12
Yes
Yes
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:wg. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (6)
5 of 13
5 of 13
5 of 13
5 of 13
4 of 13
3 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 2 )
Allele
Disease
Evidence
References
Potential Models Based on Orthology ( 2 )
Modifiers Based on Experimental Evidence ( 10 )
Allele
Disease
Interaction
References
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.
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

Please see the Physical Interaction reports below for full details
RNA-RNA
Physical Interaction
Assay
References
RNA-protein
Physical Interaction
Assay
References
protein-protein
Physical Interaction
Assay
References
Summary of Genetic Interactions
Interaction Browsers

Please look at the allele data for full details of the genetic interactions
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
suppressible
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
suppressible
suppressible
suppressible
suppressible
suppressible
External Data
Subunit Structure (UniProtKB)
Monomer; folds by intramolecular disulfide bonds (PubMed:11821428). Interacts with porcupine (por) (PubMed:11821428). Interacts with wls; in the Golgi (PubMed:18193037). Interacts with en (PubMed:1335365). Interacts with the proteoglycan Cow (heparan sulfate-bound form); this stabilizes wg and promotes its extracellular distribution (PubMed:25360738). Interacts with peg; the interaction facilitates short-range diffusion of wg (PubMed:34580289).
(UniProt, P09615 )
Linkouts
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
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
2L
Recombination map
2-26
Cytogenetic map
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
27F1-27F1
Limits computationally determined from genome sequence between P{PZ}wg02657 and P{EP}EP787EP787
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
27E5-27E5
27F1-27F2
(determined by in situ hybridisation)
28A1-28A2
(determined by in situ hybridisation)
28A1-28A3
(determined by in situ hybridisation)
Experimentally Determined Recombination Data
Notes

wg1 maps to the left of wgSp-1.

Stocks and Reagents
Stocks (862)
Genomic Clones (12)
 

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

cDNA Clones (234)
 

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)
    Other clones
    RNAi and Array Information
    Linkouts
    Antibody Information
    Laboratory Generated Antibodies
    Commercially Available Antibodies
     
    Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
    Cell Line Information
    Publicly Available Cell Lines
    Other Stable Cell Lines
     
    Other Comments

    Flies in which endogenous wg has been replaced with a membrane-tethered form of the protein are viable and produce normally patterned appendages of almost normal size (albeit with a developmental delay). In the prospective wing, prolonged wg transcription followed by memory of earlier signalling allows persistent expression of relevant target genes. It is suggested that the spread of wg protein is dispensible for patterning and growth, even though it probably contributes to increasing cell proliferation.

    gsb antagonizes wg signaling during embryonic cell fate specification and synaptic homeostasis.

    wg protein secretion requires endosome-to-Golgi retrieval of wls protein by the retromer complex.

    wg protein secretion promotes and requires retromer-dependent cycling of wls protein.

    wg signalling provides a genetic switch for the specification of leg versus tracheal fate in embryos.

    wg and its receptors, arr and fz2, are trafficked to the lysosome through the endocytic pathway in wing imaginal disc epithelial cells. In contrast to the attenuation of wg signalling in the embryonic epidermis, signalling in the wing is not attenuated by lysosomal targeting.

    wg is required for the allocation of cells to the female genital disc primordium, but is not required for allocation of cells to the male and anal primordia.

    Tethering experiments show that wg does not need to diffuse in order to pattern the mesoderm.

    wg is internalised by endocytosis and degraded in a lysosomal compartment.

    Molecular analyses suggest that the cysteine-rich domain of fz recruits wg, and bound wg interacts with the membrane portion of the receptor to initiate signalling. RNAi knockdown experiments show that fz and wg also require the receptor Arr1 to initiate arm signalling.

    wg protein undergoes a lipid modification in the endoplasmic reticulum. After modification, wg protein partitions as a membrane-anchored protein and is sorted into lipid raft detergent-insoluble microdomains at the cell surface, where it can be packaged for secretion.

    Downregulation of wg in the wing disc is essential for its development.

    1 allele of l(2)SH1281 recovered in a P-insertion screen.

    wg secreted from the head capsule organizes the peripheral specializations of the retina.

    wg is necessary for the dorsal-ventral polarization of leading edge cells early in dorsal closure, in the absence of which actin-cable assembly and actin-based cell process formation fails to occur properly in these cells.

    wg is required in the dorsal wing hinge for the establishment of axillary sclerite 3 and in the ventral wing hinge to restrict the formation of the axillary pouch.

    Wnt2 assists wg to specify the main tracheal trunk. Wnt2 affects tracheal development but apperently has no effect on the cuticle, whereas wg influences both.

    wg enhances htl expression.

    wg and htl act over sequential but interdependent phases of hindgut visceral mesoderm development. wg establishes the primordium.

    wg transcripts are localised apically in most tissues examined. This localisation is mediated by two independently acting elements in the 3' UTR. The apical localisation of the wg transcript is required for normal signalling activity of the wg protein.

    wg protein expressed in the prospective anal pads is necessary for the activation of hh in the adjacent prospective rectum in the developing hindgut. wg expression is also required for the development of the anal pads.

    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.

    nej function is necessary for ci-mediated transactivation of wg during embryogenesis.

    wg signaling has two distinct roles in tracheal development, inducing dorsal trunk fate and fusion cell fate.

    The secreted proteins encoded by hh, wg and dpp are expressed in the peripodial membrane yet they control the expression of Dl and Ser in the disc proper.

    wg and slp1/slp2 act in a common pathway in which slp1/slp2 serves as a direct target of wg signals that mediates wg effects in both ectoderm and mesoderm.

    Induction of slp1 by wg involves pan binding to multiple binding sites within a wg-responsive enhancer in the 5' region of slp1. wg signalling induces striped expression of slp1/slp2 in the mesoderm, providing striped mesodermal domains competent to respond to subsequent slp1/slp2-independent wg signals that induce somatic muscle and heart progenitors. In wg expressing ectodermal cells, slp1/slp2 is an integral component in an autocrine feedback loop of wg signalling.

    wg activates separate, yet parallel, signalling cascades that are required to promote dorsal closure and establish ventral pattern. wg is required to activate dpp expression and co-ordinate cell shape changes during dorsal closure.

    The wg gene is transcribed in narrow stripes of cells abutting the source of hh protein. These cells or their progeny are free to roam towards the anterior, away from the hh signal, whereupon wg transcription stops. The cells leaving the expression domain retain inherited wg protein in secretory vesicles, and carry it forwards over a distance of up to four cell diameters. Evidence also suggests wg protein can reach distant target cells by a second mechanism, independently of protein inheritance, possibly by restricted diffusion.

    The pnr/ush complex serves as a repressor and a transcriptional activator, respectively for wg and ush expression.

    wg is required for the appearance of tip cells in Malpighian tubules.

    N protein responds differently to binding by Dl or wg protein. The Dl signal is transduced by the N intracellular domain released from the plasma membrane, the wg signal is transduced by the N intracellular domain associated with the plasma membrane.

    fz and fz2 function as redundant receptors for wg during embryonic development. fz and fz2 function downstream of wg and upstream of arm in the wg signalling pathway.

    fz and fz2 are required during embryogenesis to maintain epidermal en and wg expression.

    wg modulates the effects of dominant negative N alleles in the developing wing of Drosophila.

    fz and fz2 are functionally redundant and act as the primary receptors for wg protein.

    wg signal transduction is abolished in virtually all cells lacking both fz and fz2 in embryos as well as in imaginal discs.

    The level and vectorial orientation of the wg concentration gradient in the notum is not important for the positioning of the dorsocentral mechanosensory bristle cluster. wg has only a permissive role on dorsocentral ac-sc expression. pnr and ush are main effectors of the regulation of wg expression in the presumptive notum.

    The en-dependent signal acts unidirectionally in the embryonic epidermis and wg activity imposes the asymmetry upon the epidermal pattern.

    wg mutant embryos have residual mirror-symmetric pattern due to an en-dependent signal specifying anterior denticle fates.

    Five EMS induced alleles have been identified in a screen for mutations affecting commissure formation in the CNS of the embryo.

    wg patterns growth in the wing primordium by modulating dm expression.

    wg interacts synergistically with Egfr to promote tergite and sternite identities in the adult abdomen, and Egfr and wg activities are opposed by dpp signalling which promotes pleural identity. wg and dpp compete directly by exerting opposite effects on cell fate. Within the tergite, the requirements of wg and Egfr function are complementary: wg is required medially, whereas Egfr is most important laterally.

    wg and dpp participate in mutual transcriptional repression in the adult abdomen.

    wg protein transport in an anterior direction generates the normal expanse of naked cuticle within the segment and movement of wg protein in a posterior direction specifies diverse denticle fates in the anterior portion of the adjacent segment in the embryo.

    Transcriptional regulation of ovo integrates inputs from the wg and Egfr pathways and control epidermis differentiation.

    wg specifies the naked fate in the embryonic epidermis at a range of up to 5 cells in the anterior direction but only in adjoining cells posteriorly.

    wg acts over a different range in the anterior and posterior directions in the embryonic epidermis. The asymmetry follows in part from differential transport or stability of wg protein: wg transport is restricted through the en domain, and at the segment boundary (in a hh-dependent manner). wg signalling represses rho expression.

    fu is required autonomously in anterior cells neighboring hh to maintain ptc and wg expression. wg is in turn maintaining en and hh expression.

    The strong genetic interactions between N and wg is at least partly due to regulation of expression of cuticle patterning genes by wg and the two forms of N.

    The levels of glycosaminoglycans (in which sgl plays a role) are rate limiting for cell-cell signalling pathways such as those of wg and hh, which mediate changes in gene expression.

    Different Wnt/Fz signals activate distinct intracellular pathways, and dsh discriminates among them by distinct domain interactions.

    High levels of fz2 protein stabilise wg protein, allowing it to reach cells far from its site of synthesis. The expression of fz2 is repressed by wg signaling, creating a gradient of decreasing wg protein stability moving toward the dorso-ventral boundary. The repression of fz2 is essential for the normal shape of the wg morphogen gradient as well as the response of cells to the wg signal. In contrast to other ligand-receptor relationships where the receptor limits diffusion of the ligand, fz2 broadens the range of wg protein action by protecting it from degradation.

    The products of the gro and pan genes interact to repress wg signalling activity.

    Activation of the N receptor in the wing disc induces strong mitotic activity. The effect of N on cell proliferation is not simply due to the upregulation of either vg or wg. On the contrary, vg and wg proteins show synergistic effects with N signaling, resulting in the stimulation of cell proliferation in imaginal discs.

    Activation of the N receptor in the wing disc induces the expression of vg and wg.

    Dorsal-ventral patterning of the eye imaginal disc develops gradually and wg plays an important role in setting up the pattern. Manipulations in wg expression shift the D-V axis of the disc. wg appears to coordinately regulate multiple events related to D-V patterning in the developing retina.

    wg is necessary and sufficient to induce dorsal expression of mirr prior to the start of differentiation and also to restrict the expression of the Ecol\lacZWR122 marker to differentiating photoreceptors near the equator.

    In wing development, wg, in conjunction with N, induces G1 and G2 arrest in separate subdomains of the zone of non-proliferating cells at the developing wing margin. The wg product induces G2 arrest in two subdomains by inducing ac and sc, which down-regulate stg. N activity creates a third domain by preventing arrest in G2 in wg-expressing cells resulting in their arrest in G1.

    The roles of N, wg and vg during the initial stages of wing development are investigated. vg is involved in the specification of the wing primordia under the combined control of N and wg signalling. Once cells are assigned to the wing fate, their development relies on a sequence of regulatory loops that involve N, wg and vg. During this process, cells that are exposed to the activity of both wg and vg will become wing blade and those that are continuously under the influence of wg alone develop as hinge. The growth of the cells in the wing blade results from a synergistic effect of the three genes N, wg and vg on the cells that have been specified as wing blade.

    A high level of dpp expression in the leg disc is required to interact with ectopic wg to induce transdetermination.

    Ectopic wg expression non-cell autonomously induces vg expression in leg discs and activated arm cell-autonomously induces vg expression, indicating that vg expression is directly activated by wg signalling.

    Segment polarity gene expression is necessary for the survival of specific rows of epidermal cells.

    wg is required for the development of the anterior protocerebral brain region in embryos.

    The dorsolateral fat body is repressed by wg and the ventral fat body needs wg for its specification. There is a balance between fat body and somatic gonadal precursor (SGP) development with tin, wg and en driving cells in the primary clusters towards SGP development and srp driving them towards fat body development.

    vn plays a permissive role in the induction of the endoderm by dpp and wg, which in turn up-regulate vn expression in the midgut mesoderm in two regions overlapping the dpp sources.

    slmb coordinates wg and dpp expression in the dorsal-ventral and anterior-posterior axes during limb development.

    The nej gene product represses pan to antagonise wg signalling.

    cad acts in hindgut development through fog, fkh and wg, but does not play a role in activating tll, hkb, byn and bowl which are also required for proper hindgut development. cad, fkh, byn and wg constitute a conserved constellation of genes that plays a required role in gastrulation and gut development.

    wg can repress its own expression via an autorepressive feedback loop that results in a change of the wg signalling profile during development.

    fz2 and not fz acts in the wg signalling pathway for wing margin development.

    A single dominant negative form of fz or fz2 can block more than one type of Wnt signalling pathway implying that truncated proteins of the Fz family lose some aspect of signaling specificity.

    CrebA wg double mutant phenotype confirms that CrebA is not involved in segment polarity.

    sgl is required for efficient wg function.

    dpp and wg act together in the regulation of furrow initiation and progression.

    dpp autoregulation and dpp-mediated inhibition of wg expression are postulated to be required for their role in eye development.

    Clonal phenotype of genes known to play compartment specific functions demonstrate the anterior/posterior patterning functions of these genes are conserved in the genital disc.

    Regulatory relationships between hh, dpp and wg in the eye are similar to those found in other imaginal discs such as the leg.

    Genetic interaction studies reveal gsb antagonises the wg signal that confers neuroblast 4-2 fate.

    Cross-regulatory relationships among hh, wg and en, as well as their initial mode of activation, in the anterior head are significantly different from those in the trunk.

    wg is essential for induction of limb primordia, specifying the most distal limb identity, dpp is not required for the induction itself.

    The specification of naked cuticle cell fate and the generation of denticle diversity by wg signaling in embryos may be generated by two distinct cellular pathways.

    sgg positively regulates dpp expression and negatively regulates wg expression in the imaginal discs, while dsh activity suppresses dpp expression and promotes wg expression in the imaginal discs.

    Activity of lbe and lbl in the epidermis is regulated by the segment polarity gene network and depends on wg signalling.

    wg and dpp act directly to provide positional information along the proximal-distal (P-D) axis of the leg. The combined activity of both signals is required in a spatially graded manner to define the distinct domains of gene expression along the P-D axis.

    ct acts to maintain margin wg expression, providing a potential explanation of the ct mutant phenotype. N, but not wg signalling, is autonomously required for ct expression. wg is required indirectly for ct expression, results suggest this requirement is due to the regulation by wg of Dl and Ser expression in cells flanking the ct and wg expression domains. Dl and Ser play a dual role in the regulation of ct and wg expression.

    Localised expression of wg in cells at the dorsal-ventral boundary of the wing can act directly at long-range to activate expression of target genes such as Dll and vg, and to control growth of the wing.

    wg signals through arm. wg can activate arm in a cell line.

    smo activity is required for transduction of hh but not wg.

    A wg response sequence (WRS) is identified in the visceral mesoderm enhancer from Ubx.

    The expression of wg and en in the adult antenna seem to be controlled by age-dependent mechanisms.

    Expression of wg and oc throughout the entire second instar eye-antennal disc confers a default fate of dorsal vertex cuticle. Activation of dpp expression in the posterior eye disc eliminates wg and oc expression, thereby permitting eye differentiation.

    Signalling by the Egfr protein antagonises signalling by the wg protein in cells of the prospective row 1-4 zone of the abdominal denticle belts in Drosophila larvae.

    Misexpression of wg in the developing eye has a potent polarizing effect on the retinal epithelium.

    hh and ptc can regulate transcription from a wg enhancer element containing ci protein binding sites by modulating the activity of ci protein.

    Casein kinase II (encoded by CkIIα and CkIIβ) phosphorylates dsh in vitro. This phosphorylation is regulated by fz2, implicating a role for Casein kinase II in wg signalling.

    wg--ptc double mutants exhibit defects in the restriction of dorsal median cells to segment boundaries and alterations in CNS and midline cell fates.

    The combined effect of N and its target genes ct and wg regulate the expression of N ligands Dl and Ser which restrict N signalling to the wing dorsoventral boundary.

    pan provides a molecular mechanism for gene control by wg signalling.

    dsh, in addition to transducing wg signal, blocks N signalling directly, explaining the inhibitory cross talk observed between the pathways.

    hh and wg play opposing roles in mesoderm segmentation.

    hh is required for the normal activation of bap and srp in anterior portions of each parasegment, whereas wg is required to suppress bap and srp expression in posterior portions.

    fz2 can function as a receptor of wg.

    The autocrine wg signal is responsible for conferring NB4-2 identity to NB5-3 in mutants.

    wg and hh signaling account for all cell types across the dorsal epidermis, where lin regulates the late expression of wg.

    Distinction between dorsal and ventral fates is maintained through mutual repression by dpp and wg.

    Expression of wg and dpp in their normal domains depends on the hh signal. Cells that are not likely to be within range of the wg or dpp signals have a different capacity to respond to hh.

    Ectopic expression of wg during eye development causes lack of interommatidial bristles; the wg signal transduction machinery found in the embryo and wing also functions in the eye. In the absence of N, wg signaling appears to occur normally, arguing against a direct role for N in wg signaling.

    Ectodermal and mesodermal Dr expression depend on wg and hh.

    hh signalling in the germarium involves ptc and ci, but not wg or dpp.

    wg accumulation at the DV border in the wing sets, by repression, the gap between the two patches of ara-caup expression.

    hh, wg and dpp are required for the establishment of signaling centres that coordinate morphogenesis in the hindgut epithelium.

    Activation of hh, wg and dpp in the developing hindgut and foregut requires fkh. hh and wg activities in the gut epithelial cells are required for the expression of the homeobox gene bap in the ensheathing visceral mesoderm.

    Complementary and mutually exclusive activities of dpp and wg organise axial patterning during leg development.

    dpp participates with wg to induce transdetermination. The dose-dependent effects of wg support a model in which wg functions as a morphogen in imaginal discs.

    Ectopic expression of wg is all discs reveals a dose-dependent effect of wg on patterning and transdetermination that is mediated by modulation of dpp expression.

    por is involved in wg processing. por acts nonautonomously in wg patterning.

    No evidence has been found to suggest that wg regulates vg expression in the developing wing blade.

    A combination of hh and wg is required to specify the most posterior fates of the A compartment. wg could play the role of the morphogen, at least within part of the segmental pattern.

    Both dpp and wg are required for the nuclear localization of exd protein in the endoderm. Ectopic expression of dpp and wg expands the domain of nuclear exd. The nuclear import of exd product correlates with the transcription of an exd-dependent reporter gene in the endoderm.

    Four segment polarity genes, hh, wg, gsb and en all function in concert to determine the formation and specifications of three hh-dependent eg-neuroblasts (6-4, 7-3 and 2-4).

    dpp can act both synergistically with and antagonistically to wg in leg disc patterning.

    Overexpression of dpp has little effect on dorsal leg patterning, limited effects on wing patterning but substantial dose-dependent effects on anterior-ventral leg patterning.

    A hierarchical relationship between N, wg and vg patterns the dorsoventral axis of the wing.

    Expression of wg and vg in the wing margin are direct and parallel responses to the activation of N. wg is not required for the activation of vg, wg activation does not depend on vg function at the dorsoventral boundary. Expression of vg in the wing pouch depends on wg activity, suggesting that a secondary function of vg is to mediate the long-range effects of secreted wg protein in the wing pouch. wg and N cooperate to activate expression of ct, suggesting the wg and N pathways interact synergistically in the wing imaginal disc.

    The 'spade-flag' mutant phenotype caused by wgspd-fg results from the loss of wg expression in the wing hinge causing decreased proliferation in the hinge region. Ectopic activation of the wg pathway causes overproliferation in the hinge region of the wing.

    'Sternopleural' is a regulatory allele of wg, wgSp-1. Genetically Sp maps to the 3' regulatory region of wg and is lethal when heterozygous with two wg alleles that affect the 3' regulatory region. In addition a mutation molecularly mapped to the 3' regulatory region exhibits a mild Sp phenotype. Interallelic complementation between wg alleles can best be explained by transvection.

    wg protein has two distinct functions in wing formation, a primary role in specification of the wing primordium and a secondary role as a mediator of the growth and patterning activities of the dorso-ventral compartment boundary. Both wg and vg proteins are required to promote the growth of the wing but only after the wing field has been established.

    Ventral expression of wg is a consequence of activity of the dorso-ventral system and therefore acts downstream of the N and Ser proteins. Both wg and vg proteins are targets for activation by the dorso-ventral system.

    wg is necessary for heart formation.

    wg overexpression in the early gastrula causes an expansion of the cardiac mesoderm. wg overexpression wg restores the heart deficit of hh mutant embryos. dsh overexpression causes hypertrophy of heart precursors and rescues the heart and CNS deficits of wg mutants.

    dpp restricts the domain of wg during limb patterning.

    By repressing wg expression in the leg, dpp signalling limits the response to high levels of wg and dpp to the site of distal outgrowth.

    wg acts as an inductive signal that influences the expression of nau in the ventral mesoderm.

    Secreted wg protein can bind to glycosaminoglycans with high affinity, and interaction with glycosaminoglycans can promote wg signal transduction.

    hh and wg specify distinct head capsule region identities.

    During eye-antennal disc development hh and wg expression initially overlap, but subsequently segregate. This regional segregation is critical to head specification and is regulated by oc.

    The boundary between wg-expressing cells of the wing margin and the adjacent proneural cells, which give rise to the margin sensory bristles, arises in part by a mechanism of "self-refinement" where wg protein represses wg expression in adjacent cells. Cells unable to receive the wg signal do not resolve the boundary between wg-expressing and proneural cells, consistent with the hypothesis that wg inhibits N. arm is not required for wg self refinement at the wing margin.

    Overexpression of full length or truncated shg phenocopies wg. This is achieved by titrating arm away from a signalling function in the wg pathway.

    wg is expressed in the presumptive tissues of the adult abdomen during their development and is involved in several patterning processes. The requirements for wg in bristle and cuticle formation are separable. The timing of appearance of bristle precursors correlates with the sensitive period for wg, suggesting wg is required for the determination of sensory organ precursor cells.

    Loss of dpp signalling leads to ectopic wg gene expression and loss of wg signalling leads to ectopic dpp gene expression. Increased dpp signalling represses wg transcription and increased wg signalling represses dpp transcription.

    wg transcription is repressed by dpp in the developing eye.

    sgg, dsh and arm function to transmit the wg signal in the midgut in the same way as they do in the epidermis. The wg signal transduction pathway acts in all three germ layers, the ectoderm, mesoderm and endoderm.

    dsh and arm mutations mimic all effects of a wg mutation. sgg mutations emulate the effects of high ubiquitous wg in the midgut.

    wg can act directly and at long range as a gradient morphogen during normal development.

    Wild-type wg acts at long range, up-regulating the transcription of particular target genes as a function of concentration and distance from secreting cells. Tethered wg has only short range effects on the transcription of target genes.

    Clones of dsh mutant tissue at the wing margin can block the response to wg.

    wg is expressed in a narrow stripe at the wing margin, dsh is required in the cells responding to wg.The wg signal can traverse several cell diameters of mutant tissue to reach responsive wing cells. Overexpression of dsh potentiates the response to wg, bristle formation is induced large distances from the site of wg expression. wg can signal over a large distance in a sensitized background.

    wg and dpp are secreted by ectodermal cells and the proteins are required for the differentiation of the underlying mesodermal cells and for the segregation of the somatic and visceral muscles.

    wg acts directly in the mesoderm to ensure the formation of slou-expressing founder cells.

    wg gene product can signal across germ layers so wg from the ectoderm could constitute an inductive signal for the initiation of the development of a subset of somatic muscles.

    wg signalling may involve complex interactions between the wg ligand and its cell surface receptor molecule(s) and that some of this complexity resides within the wg ligand itself.

    The wg signalling pathway is involved in mesodermal pattern formation.

    wg function is not required for overgrowth in l(2)gd1 leg discs but is required for disc duplication.

    wg is a ventral signal during wing development.

    Induction of vg requires the combined activities of Ser, wg and N. The combination of Ser (dorsal signal) and wg (ventral signal) at the dorso-ventral interface activates the N receptor and leads to vg expression.

    wg mediates the organising activity of the wing dorsal/ventral compartment boundary. wg activity on either side of the boundary is sufficient to support development of the wing blade.

    Activation of N, mediated through Su(H), is sufficient to direct wg expression in the wing dorsal/ventral boundary. Localised expression of Ser in the dorsal wing cells provides the signal to activate wg.

    wg and Wnt4 achieve opposite, but complementary functions in intrasegmental cell patterning of the embryonic ectoderm.

    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.

    The wg and N signalling pathways interact to define the range of lateral inhibition within the SNS anlage, causing the selection of the three ac expressing cells instead of one.

    Invagination centers within the somatogastric nervous system anlage are positioned by N-mediated signaling which is spatially controlled though wg.

    wg function in the midgut is absolutely required for copper cell development.

    High levels of wg repress copper cell development but allow the development of large flat cells and ubiquitous levels of wg can produce ectopic copper cells.

    Pka-C1 and hh have antagonistic effects on a common substrate which regulates transcription of dpp and wg.

    Pka-C1 is essential during limb development to prevent inappropriate dpp and wg expression. A constitutively active form of Mmus\Pkaca, can prevent inappropriate dpp and wg expression but does not interfere with their normal induction by hh. The basal activity of Pka-C1 imposes a block on the transcription of dpp and wg and hh exerts its organizing influence by alleviating the block.

    Ectopic expression of wg can elicit transdetermination in all ventral appendages, including structures that arise from labial, antennal, maxillary palpus and genital primordia, as well as in that of all three legs. The locations of the transdeterminations all map to dorsal regions of the appropriate imaginal disc. wg does not induce transdetermination in dorsal appendages, wing, haltere or humerus.

    wg and dpp are involved in the organization of the visual ganglia.

    wg expression is necessary for the induction and maintenance of dpp expression in adjacent cells during optic lobe development.

    Expression of wg is ventral and expression of vertebrate wg homologs is dorsal.

    The development of eve cells (cells from parasegments 4-12 that give rise to the pericardial cells of the heart) depends on at least wg and hh. Two classes of mosaic clones demonstrate that wg protein in either ectoderm or mesoderm is sufficient for the development of eve cells in the mesoderm and the patterning the mesoderm.

    Uniform expression of wg in the mesoderm alone is sufficient to rescue the repeated clusters of eve expressing cells.

    Pka-C1 is an integral component of the mechanism that restricts the expression of dpp, ptc and wg in imaginal discs.

    Germ line transformation demonstrates that the 3' untranslated region of wg is sufficient for apical localisation of the wg transcript.

    wg and ptc are negative regulators of the morphogenetic furrow and influence tissue polarity in the developing compound eye.

    por is required for direct wg autoregulation and is epistatic to sgg in the regulation of wg. por is required for autoregulation by exogenous wg.

    Ectopic expression of wg induces transdetermination of dorsal leg imaginal disc cells to ventral wing cells, this transdetermination is very similar to the leg-to-wing switch that occurs after leg disc culture.

    ci is epistatic to ptc in the maintenance of wg expression and the formation of naked cuticle.

    wg expression is regulated by odd and prd. odd represses wg expression, prd restricts the domain of expression of odd.

    The transition from inter-dependent wg and en expression to wg autoregulation may involve several segment polarity genes.

    hh, wg and mys are required for epithelial morphogenesis during proventriculus organ development.

    The en-hh-ptc regulatory loop that is responsible for segmental expression of wg in the embryo is reused in imaginal disks to create a stripe of dpp expression along the A/P compartment boundary.

    wg performs at least two roles in the patterning of the adult epidermis, a neurogenic role and a differentiation role. Histoblasts are competent to respond to the neurogenic signal at an earlier stage than the differentiation signal, though the two phases overlap. Loss and gain of function experiments suggests that bristles are determined, in response to wg, at a very early stage of abdominal development, well before they have reached their final position.

    wg acts to prevent dpp present at the dorsal and ventral margins in the regions that will form head cuticle from initiating a wave of photoreceptor development. wg and dpp interact to define the region in which the morphogenetic furrow can initiate.

    Ectopic wg can inhibit the propagation of normal photoreceptor development.

    The wg product has two functions in the leg disc. In its specification of ventral fate, the wg product does not act as a morphogen. wg inhibits the commitment of dorsal cells toward a determined state and influences the regulation of proliferation.

    The effect of wg expression on the proximo-distal axis is independent of its function in dorsal-ventral specification.

    wg function is specifically required for heart development. The developmental requirement of wg for cardiac organogenesis is distinct from its function in segmentation and neuroblast identity.

    dsh is not merely a permissive factor in wg signalling but encodes a novel signal transduction molecule which may function between the wg receptor and more downstream signaling molecules.

    Maintenance of restricted wg transcription during late gastrulation requires en-independent wg autoregulatory activity. The spatial localization of the wg product is important in the asymmetric patterning of epidermal cell fates.

    hh regulates wg expression.

    opa activity is essential for the appropriate level and timing of en and wg expression in all parasegments, but does not determine their restricted spatial domains of expression.

    The ptc membrane-bound product partially co-localizes with the wg product during embryogenesis, suggesting the interaction of ptc protein with elements of the reception complex of wg.

    Genetic interaction and phenotypic analysis suggest that the product of wg is a ligand for N during pattern formation in the embryo and the imaginal discs.

    arm, dsh and sgg encode elements of a unique wg signalling pathway that is used several times throughout development.

    wg acts through inactivation of the sgg protein kinase to specify ventral cell fate in the leg.

    Combined action of wg-expressing cells in the leg ventral anterior compartment and dpp-expressing cells in the leg dorsal-anterior compartment activates expression of Dll.

    The maintenance of wg expression by the hh signal is limited to early development.

    N is necessary for the implementation of the wg signal in specifying normal wing development.

    Direct wg autoregulation differs from wg signalling to adjacent cells in the importance of fu, smo and ci relative to sgg and arm. Early wg autoregulation during the hh-dependent stage differs from later wg autoregulation.

    The assembly of the target structure for ingrowing retinal axons involves cell-cell interactions mediated by the wg product. wg and dpp are required for target field neurons to adopt their proper fates and to send axons into the developing target structure.

    wg product made in the mesoderm can sustain en expression in the ectoderm.

    Antp, Ubx, abd-A, dpp and wg are required for proper tsh expression.

    wg acts through dsh and arm to affect the expression of en and cuticle differentiation.

    Comparisons of early development to that in other insects have revealed conservation of some aspects of development, as well as differences that may explain variations in early patterning events.

    Double mutant analysis demonstrates that arm's role in wg signalling is direct; arm functions downstream of wg and sgg.

    wg is a target gene of the homeotic selector genes and is regulated by exd.

    Repeated pulses of ptc lead to transcriptional repression of the segment polarity genes wg and gsb in the embryonic trunk and a wg like phenotype in the larval cuticle.

    dsh and por act upstream of sgg, and arm acts downstream of sgg in the wg signalling pathway.

    dsh is required during wg signalling to establish both cell polarity and cell identity.

    Mosaic analysis suggests that wg-expressing cells sustain en expression only in adjoining cells. The autoregulatory effect of wg on its own expression is also short range.

    Cell culture assay of wg and arm gene expression demonstrates that the wg protein does not affect the rate of arm protein synthesis but presence of the wg protein causes increased stability of an otherwise rapidly decaying arm protein. wg protein from the co-culturing donor cells, in the extracellular matrix and soluble medium from donor cells also increases the levels of arm protein demonstrating that wg can act as a soluble extra cellular signalling molecule.

    Wild type activity of five segment polarity genes, wg, ptc, en, nkd and hh, can account for most of the ventral pattern elements in the embryo. wg is required for naked cuticle. wg generates the diversity of cell types within the segment but each specific cell identity depends on the activity of ptc, en, nkd and hh.

    Maintenance of en expression does not always require wg activity.

    The eve product represses wg expression.

    The presumptive tips of both the leg and wing are characterized by the close association of cells expressing wg, dpp and al. Ectopic expression of wg can induce ectopic al expression, but only in regions with high levels of dpp.

    Ectopic expression of wg can induce a duplication of the proximodistal axis, but only in regions with high levels of dpp.

    Using a temperature sensitive allele it was shown that the wg expressing cells in the neuro-ectoderm generate normal neuroblasts after wg inactivation, but the anterior and posterior neurectoderm requires wg nonautonomously for subsequent neuroblast determination and formation.

    In the embryo thoracic imaginal primordia are allocated in response to signals from wg and dpp. wg is required for allocation and for initiation of Dll expression that marks the allocation process.

    Temperature shift experiments and the wg expression pattern in sectors in imaginal discs provide references for a polar coordinate system homologous to that postulated in a model for regeneration in insect and vertebrate wings (FBrf0029389, FBrf0037332, Bryant, Trends Genet. 2:153 ).

    wg functions as an extracellular signal maintaining en expression.

    Transcriptional control of both ptc and wg by hh is mediated by the same signal transduction pathway. fu and ci are required for normal wg transcription, acting downstream of ptc to regulate wg transcription. cos negatively regulates ptc and wg transcription.

    Segment polarity mutations cause stripes of abnormal patterning within sectors of the leg disc, which may be mediated by regional perturbations in growth.

    wg activity is required only indirectly for wg expression at the parasegment boundary.

    The ptc and hh genes encode components of a signal transduction pathway that regulate the expression of wg transcription following its activation by pair rule genes, but most other aspects of wg expression are independent of ptc and hh. Expression of wg in the absence of ptc depends on hh. Absence of ptc activity can result in de novo activation of wg after gastrulation.

    Competence of cells to express wg is independent of their ability to receive the hh signal. wg activation requires the function of fu, this suggests that the putative hh signal is transduced by the serine/threonine kinase that fu encodes.

    wg plays a role in the regulation of run mRNA expression.

    The main function of gsb is the maintenance of wg expression by a wg-gooseberry autoregulatory loop after 6 hours of development. A temporal asymmetry in the regulatory interactions between wg, en and the gsb explains the difference in phenotypes between the mutants.

    wg may regulate the activity of the achaete-scute complex in proneural clusters. wg function is also required at the presumptive wing margin and is a necessary precondition for the change in proliferation pattern at the wing margin.

    wg does not act as a local instructive signal or a morphogen. The wg protein may maintain and seal parasegmental borders, isolating abutting segmental gradients.

    Overexpression of wg protein does not prevent parasegmental, restricted en expression.

    wg-expressing cells normally give rise to the most ventral portion of the leg. wg protein organises leg pattern along the dorsoventral axis by conferring ventral positional information in the disc. Varying levels of wg activity specify varying portions of dorsoventral pattern.

    Positive autoregulation of Ubx is at least partly indirect and is mediated by the extracellular signal molecules encoded by dpp and wg : indirect autoregulatory mechanisms may be used to ensure coordinate maintenance of selector gene activity in groups of cells.

    The upstream sequence of Ubx contains separate response elements for dpp and wg which function independently, though both positive and negative interactions take place between them.

    The wg product is required to restrict the expression of ap gene to dorsal cells in the developing wing and to promote the expression of vg and sd that demarcate the wing primordia.

    Wild type wg alleles transfected into Drosophila tissue culture cells display wg protein on the cell surface and in the extracellular surface, whereas mutant proteins appear not to be secreted. Cells from embryos mutant for por show a retention of the wg product, suggesting that por provides an accessory function for wg protein secretion or transport.

    Choice of cell fate made by en expressing cells in embryonic parasegments is mediated by wg, in a function distinct from its early role in maintaining en expression. en expressing cells respond differently to wg at different stages of development:early wg stabilizes the subdivision of the body axis by maintaining en expression, whereas later input generates cell-type diversity.

    The effects of wg mutants on hh expression appear earlier than the effects of en mutants on hh expression.

    The wg gene is involved both in controlling the segmentation pattern of embryos by affecting the posteriormost cells of each parasegment and in controlling the imaginal disk pattern of the meso- and meta-thoracic segments that develop into wing, haltere and notum in pupae and adults. The temperature-sensitive period for wgl-12, a temperature-sensitive allele, lies between gastrulation and the beginning of dorsal closure (11 hours after egg laying at 25oC).

    Mutant alleles of wg may be viable, showing a visible phenotype involving loss of wings and/or halteres, or they may be homozygous lethal (usually as embryos but sometimes as pupae). Low temperature fails to rescue heteroallelic combinations of wg1 or wgl-14 with the temperature-sensitive allele wgl-12 after the larval stages.

    On basis of ectopic eve expression experiments, it has been suggested that eve is a potent and direct repressor of wg.

    Derepression of en in Pc group mutants is not acting through wg.

    Most cells in embryo can interpret wg signal, therefore tight control of wg expression pattern is essential for normal development.

    Sequence analysis of Wnt genes is performed in several species to determine the ancestral lineage of the gene family.

    Genetic epistasis experiments indicate that wg signalling operates by inactivating the sgg repression of en autoregulation. sgg is epistatic to wg.

    Cell division in the Malpighian tubules depends on normal wg expression.

    Over-expression of wg results in supernumerary cells in the tubules.

    In the absence of ptc function, wg expression, which is normally en-dependent, no longer requires en.

    wg and en function in patterning the larval epidermis.

    wg protein is secreted in the embryo and is taken up by neighbouring cells. The protein can be found two or three cell diameters away from the cells in which it is synthesised.

    wg cannot completely rescue the ptc phenotype when in double mutant combinations.

    The role of ptc in positional signalling is permissive rather than instructive, its activity is required to suppress wg transcription in cells predisposed to express wg. These cells receive an extrinsic signal, encoded by hh, that antagonises the repressive activity of ptc. The ptc protein may be the receptor for the hh signal.

    Mutations in zygotic polarity gene wg do not interact with RpII140wimp.

    Mutations in arm and wg have indistinguishable embryonic consequences.

    Clonal tissue with reduced levels of arm activity will only survive in regions furthest from regions of high levels of wg RNA.

    The notum duplications seen in apID and apXa resemble notum duplications produced by wg mutations. Double heterozygotes of wg and ap demonstrate that the two gene products are interact.

    wg plays an important role in defining the positions in which leg primordia will develop along the antero-posterior axis of the embryo.

    wg is essential for maintaining the normal pattern of ptc expression.

    Ubx, abd-A, dpp, wg and lab are involved in the induction process between the visceral mesoderm and the gut epithelium in the embryo.

    Ubx, abd-A, dpp, wg and lab have interacting gene products. abd-A function is necessary for wg expression.

    abd-A function is required for expression of wg in the visceral mesoderm cells of the anterior midgut posterior to dpp expressing cells.

    wg regulates accumulation of arm by post-transcriptional control, por and dsh are also required for this effect.

    wg has a specific role in the control of cell fates during neurogenesis.

    In the nervous system, a single neuron, RP2, is missing; other neurons in the lineage are normal.

    Genetic analysis demonstrates that wg is dispensable for efficient homeotic gene expression in the visceral mesoderm.

    Dorsal abnormalities in mutant embryos are more extreme than ventral ones, the dorsal cuticle being greatly reduced and covered by fine hairs. The adult wingless phenotype shows incomplete penetrance and variable expressivity and is affected by the ability of the wingless gene to function during the larval period. Lethal as well as viable wg alleles are not cell-autonomous in adult mosaics.

    wg is believed to control segment organization through an intercellular signaling mechanism.

    The wg gene is involved in controlling the segmentation pattern of embryos by affecting the posteriormost cells of each parasegment. wg is believed to control segment organization through an intercellular signaling mechanism.

    In embryonic lethal alleles, each segment shows a mirror-image duplication of the denticle bands at the expense of naked cuticle so that a continuous sheet of denticles (instead of repeated denticle bands) is produced.

    Mutant embryos lack head structures and filzkorper.

    The wg gene is believed to control segment organization through an intercellular signaling mechanism.

    Genetic mosaics were used to determine that wg is not autonomous at the level of the single cell.

    Lethal as well as viable wg alleles are not cell-autonomous in adult mosaics.

    In pupal lethal and adult viable alleles, the ready-to-emerge pupae and the adults lack one or both wings and/or halteres and there is a corresponding duplication of the meso- and metanota.

    In pupal lethal and adult viable alleles, the ready-to-emerge pupae and the adults lack one or both wings and/or halteres and there is a corresponding duplication of the meso- and metanota. Lethal as well as viable wg alleles are not cell-autonomous in adult mosaics.

    The wg gene is involved both in controlling the imaginal disk pattern of the meso- and meta-thoracic segments that develop into wing, haltere and notum in pupae and adults.

    Relationship to Other Genes
    Source for database merge of

    Source for merge of: wg Gla

    Source for merge of: wg l(2)SH1281

    Additional comments

    The Sternopleural and spade mutations interact with wg alleles. spade is a defect in a cis-regulatory region necessary for correct imaginal disc expression of the wg gene, but Sternopleural defines a distinct function. The developmental events that are normally explained by wg activity only might be better explained by a consortium of molecules encoded by genes that map very close to wg, that work together to pattern various tissues.

    wg is functionally analogous to mouse Wnt-1 in mammary cell transformation assays, causing transformation via a paracrine mechanism.

    Pairwise complementation analysis of wg1, wgl-8, wgH and an allele of wg reveals a complex complementation pattern.

    Nomenclature History
    Source for database identify of

    Source for identity of: wg CG4889

    Nomenclature comments
    Etymology

    The name "wingless" refers to the lack of wings observed in mutants.

    Synonyms and Secondary IDs (41)
    Reported As
    Symbol Synonym
    Dint-1
    Wg
    (Akai et al., 2025, Dias and Dilão, 2025, Farmer et al., 2025, Hingole et al., 2025, Hounsell and Fan, 2025, Park et al., 2025, Vega-Cuesta et al., 2025, Fischer et al., 2024, Goins et al., 2024, Jiménez-Jiménez et al., 2024, Kumar et al., 2024, Monticelli et al., 2024, Sachan et al., 2024, Sun et al., 2024, Wang et al., 2024, Bharti et al., 2023, Cabrita and Martinho, 2023, Chafino et al., 2023, Chen et al., 2023, Ho et al., 2023, Iyer et al., 2023, Koh et al., 2023, Sen, 2023, Baonza et al., 2022, Colon-Plaza and Su, 2022, Daly et al., 2022, Enomoto and Igaki, 2022, Hale et al., 2022, Jullien et al., 2022, Mesrouze et al., 2022, Neophytou and Pitsouli, 2022, Schember and Halfon, 2022, Zhang and Edgar, 2022, Akai et al., 2021, Al Hayek et al., 2021, Boukhatmi, 2021, Charlton-Perkins et al., 2021, Costa-Rodrigues et al., 2021, Destalminil-Letourneau et al., 2021, Dye et al., 2021, Emmons-Bell and Hariharan, 2021, Fan et al., 2021, Frankenreiter et al., 2021, García-López et al., 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Muñoz-Descalzo et al., 2010, Oros et al., 2010, Quijano et al., 2010, Reim and Frasch, 2010, Ren et al., 2010, Rhiner et al., 2010, Sawa, 2010, Silhankova et al., 2010, Sinenko et al., 2010, Swaminathan et al., 2010, Terriente-Félix et al., 2010, Wang and Hou, 2010, Wu and Johnston, 2010, Yagi et al., 2010, Zimmerman et al., 2010, Akbar et al., 2009, Benítez et al., 2009, Cruz et al., 2009, Firth and Baker, 2009, Glavic et al., 2009, Kategaya et al., 2009, Korkut et al., 2009, McKay et al., 2009, Miller et al., 2009, Nagaraj and Banerjee, 2009, Smith-Bolton et al., 2009, Svendsen et al., 2009, Wright and Tjian, 2009, Zhang et al., 2009, Acar et al., 2008, Bachmann et al., 2008, Baena-Lopez et al., 2008, Bejarano et al., 2008, Bornemann et al., 2008, Buechling et al., 2008, Canela-Xandri et al., 2008, Carrera et al., 2008, Chang et al., 2008, Chang et al., 2008, Chang et al., 2008, Chen et al., 2008, del Alamo and Mlodzik, 2008, Eid et al., 2008, Estella et al., 2008, Fan and Bergmann, 2008, Fan and Bergmann, 2008, Franch-Marro et al., 2008, Friggi-Grelin et al., 2008, Gallet et al., 2008, Herranz et al., 2008, Kaspar et al., 2008, Kennell et al., 2008, Leatherman and DiNardo, 2008, Lin et al., 2008, Miura et al., 2008, Okajima et al., 2008, Schwank et al., 2008, Smith-Bolton et al., 2008, Somorjai and Martinez-Arias, 2008, Tien et al., 2008, Ueyama et al., 2008, Vaccari et al., 2008, Wang et al., 2008, Wilkin et al., 2008, Williams et al., 2008, Zimmerman et al., 2008, Bartscherer et al., 2007, Bhambhani et al., 2007, Bhat, 2007, Buceta et al., 2007, Chan et al., 2007, Chaudhary and Strutt, 2007, Coudreuse and Korswagen, 2007, Estella and Mann, 2007, Estella et al., 2007, Gajewski et al., 2007, Garcia et al., 2007, Halachmi et al., 2007, Kennell and Cadigan, 2007, Kicheva et al., 2007, Kuranaga and Miura, 2007, Lan et al., 2007, Luo et al., 2007, Luque and Milan, 2007, Mishra and Nigam, 2007, Parker et al., 2007, Perrimon and Mathey-Prevot, 2007, Pfleger et al., 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Eldar and Barkai, 2005, Häcker et al., 2005, Hayward et al., 2005, Jordan et al., 2005, Kirkbride, 2005, Mehlen et al., 2005, Menon et al., 2005, Milan et al., 2005, Mok et al., 2005, Parker and Cadigan, 2005, Staedeli and Basler, 2005, Strutt and Strutt, 2005, Takada et al., 2005, Wu and Nusse, 2005, Cong et al., 2004, Dominguez et al., 2004, Fanto and McNeill, 2004, Huang and Klein, 2004, Kamimura et al., 2004, Matsubayashi et al., 2004, Voas and Rebay, 2004, Chinnadurai, 2003, Schweizer and Varmus, 2003, Cox and Baylies, 2002, Pandur et al., 2002, Petit et al., 2002, Bhandari and Shashidhara, 2001, Merabet et al., 2001, Saller and Bienz, 2001, Rulifson et al., 2000, Selleck et al., 2000, Magie et al., 1999, Weinmaster, 1997)
    fg
    l(2)SH1281
    l(2)SH2 1281
    l(2)rO727
    spd
    wg
    (Brutscher et al., 2025, Klemm et al., 2025, Liu et al., 2025, Ricolo et al., 2025, Slack, 2025, Alaraby et al., 2024, Baker, 2024, Balakireva et al., 2024, Breuer et al., 2024, Chang et al., 2024, Collins et al., 2024, Cruz et al., 2024, Cui et al., 2024, Datta and Bangi, 2024, Ewen-Campen and Perrimon, 2024, Jang et al., 2024, Janzen et al., 2024, Ju et al., 2024, Li et al., 2024, Melde et al., 2024, Puli et al., 2024, Raicu et al., 2024, Sharma and Chaudhary, 2024, Tan et al., 2024, Zhang et al., 2024, Anand et al., 2023, Attrill, 2023, Chan et al., 2023, Chimata et al., 2023, Floc'hlay et al., 2023, Golubev et al., 2023, Guss et al., 2023, Huang et al., 2023, Molina-Gil et al., 2023, Nandy and Roy, 2023, Neophytou et al., 2023, Petsakou et al., 2023, Posnien et al., 2023, Rosales-Vega et al., 2023, Titus et al., 2023, Voutyraki et al., 2023, Yamada et al., 2023, Beaven and Denholm, 2022, Catalani et al., 2022, Chen et al., 2022, DeBardlabon et al., 2022, Deshpande et al., 2022, 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    Name Synonyms
    Bristled
    Complementation group I
    Glazed
    Wingless
    (Joy et al., 2024, Matamoro-Vidal et al., 2024, Alvarez-Rodrigo et al., 2023, Khan et al., 2023, Markus et al., 2023, Cabana and Lussier, 2022, Kimble and Nüsslein-Volhard, 2022, Weina et al., 2022, Hayes and Melrose, 2021, Mishra et al., 2021, Morin-Poulard et al., 2021, Hayashi et al., 2020, Khadilkar et al., 2020, McGough et al., 2020, Muralidharan, 2020, Palm and Rodenfels, 2020, Song and Zhou, 2020, Wang and Dahmann, 2020, Gahr et al., 2019, Xu et al., 2019, Yadav and Eleftherianos, 2019, Alfred and Vaccari, 2018, Schwartz and Rhiner, 2018, Ahmad, 2017, Amourda and Saunders, 2017, Arnés et al., 2017, Beer and Wehman, 2017, Chen et al., 2016, Jenny and Basler, 2016, Nakato and Li, 2016, Ravisankar et al., 2016, Tian et al., 2016, Wang et al., 2016, Bieli et al., 2015, Diop and Bodmer, 2015, Hall and Verheyen, 2015, Palmer et al., 2015, Simon and Guerrero, 2015, Surabhi et al., 2015, Svendsen et al., 2015, Vlisidou and Wood, 2015, Zhang et al., 2015, Amoyel and Bach, 2014, Gunage et al., 2014, Halstead et al., 2014, Jenny and Basler, 2014, Jones and Srivastava, 2014, Kux and Pitsouli, 2014, Mishra et al., 2014, Ocorr et al., 2014, Oliveira et al., 2014, Ayyaz and Jasper, 2013, Beckett et al., 2013, Herrera et al., 2013, Hirabayashi et al., 2013, Hombría and Sotillos, 2013, Kleinschmit et al., 2013, Lawrence and Casal, 2013, Levayer and Moreno, 2013, Marques-Pita and Rocha, 2013, Morin-Poulard et al., 2013, Muha and Müller, 2013, Sasamura et al., 2013, Solis et al., 2013, van de Hoef et al., 2013, Yamamoto-Hino and Goto, 2013, Zhang et al., 2013, Dahal et al., 2012, Dani et al., 2012, Koles and Budnik, 2012, Kunttas-Tatli et al., 2012, Kuwamura et al., 2012, Mirth and Shingleton, 2012, Niehrs, 2012, Pirooznia et al., 2012, Sagner et al., 2012, Troost and Klein, 2012, Tursun, 2012, Wong et al., 2012, Zhou et al., 2012, Buechling et al., 2011, Dahmann et al., 2011, Fiedler et al., 2011, Hamaratoglu et al., 2011, Harterink et al., 2011, Leonardi et al., 2011, Momota et al., 2011, Monier et al., 2011, Port et al., 2011, Rodriguez, 2011, Singh et al., 2011, Vincent et al., 2011, Vorwald-Denholtz and De Robertis, 2011, Wang et al., 2011, Wolpert, 2011, You et al., 2011, Zhu, 2011, Beam and Moberg, 2010, Bejarano et al., 2010, Fiuza et al., 2010, Lin et al., 2010, Marie et al., 2010, Maurel-Zaffran et al., 2010, Metcalfe et al., 2010, Mohan et al., 2010, Mosca and Schwarz, 2010, Muñoz-Descalzo et al., 2010, Portela et al., 2010, Rhiner et al., 2010, Terriente-Félix et al., 2010, Varelas et al., 2010, Wang and Hou, 2010, Wu and Johnston, 2010, Yan et al., 2010, Zimmerman et al., 2010, Akbar et al., 2009, Chen and Rasmuson-Lestander, 2009, Cruz et al., 2009, Davidson et al., 2009, Eivers et al., 2009, Kategaya et al., 2009, Mao et al., 2009, Miller et al., 2009, Mirth et al., 2009, Svendsen et al., 2009, Widmann and Dahmann, 2009, Yan et al., 2009, Aoki et al., 2008, Baudot et al., 2008, Bejarano et al., 2008, Blauwkamp et al., 2008, Bornemann et al., 2008, Canela-Xandri et al., 2008, Chang et al., 2008, Chang et al., 2008, Dougherty et al., 2008, Duong et al., 2008, Estella et al., 2008, Fan and Bergmann, 2008, Legent et al., 2008, Lin et al., 2008, Lin et al., 2008, McClure and Schubiger, 2008, Pancratov et al., 2008, Sato et al., 2008, Smith-Bolton et al., 2008, Somorjai and Martinez-Arias, 2008, Song et al., 2008, Takacs et al., 2008, Tran et al., 2008, Vaccari et al., 2008, Vied and Kalderon, 2008, Williams et al., 2008, Wu et al., 2008, Zeng et al., 2008, Bartscherer et al., 2007, Bhat, 2007, Buceta et al., 2007, Bulanin and Orenic, 2007, de la Roche and Bienz, 2007, Franch-Marro et al., 2007, Gajewski et al., 2007, Galindo et al., 2007, Herranz et al., 2007, Kerszberg and Wolpert, 2007, Lai et al., 2007, Manak et al., 2007, Mann et al., 2007, Payre et al., 2007, Pellock et al., 2007, Reig et al., 2007, Schwartz and Pirrotta, 2007, Sevrioukov et al., 2007, Soller et al., 2007, Theisen et al., 2007, Tountas and Fortini, 2007, Tsai et al., 2007, Albrecht et al., 2006, Baehrecke, 2006, Beckett and Baylies, 2006, Bossing and Brand, 2006, Carmena et al., 2006, Colosimo and Tolwinski, 2006, Gonzalez et al., 2006, He and Axelrod, 2006, Hufnagel et al., 2006, Jaiswal et al., 2006, Ma et al., 2006, Maqbool et al., 2006, Nystul and Spradling, 2006, Olson, 2006, Philippakis et al., 2006, Price et al., 2006, Quaiser, 2006, Rusten et al., 2006, Singh et al., 2006, Tolhuis et al., 2006, Veitia, 2006, Wendler et al., 2006, Yasunaga et al., 2006, Dawber et al., 2005, Hoehne et al., 2005, Kirkbride, 2005, Marques, 2005, Mehlen et al., 2005, Milan et al., 2005, Mok et al., 2005, Strigini, 2005, Strutt and Strutt, 2005, Fanto and McNeill, 2004, Huang and Klein, 2004, Kamimura et al., 2004, Matsubayashi et al., 2004, Papadopoulou et al., 2004, Voas and Rebay, 2004, Fernandez-Garza and Couso, 2003, Gonzalez-Gaitan and Jackle, 2000, Wesley, 1999)
    flag
    lethal (2) SH1281
    wingless
    (Karasawa and Koshikawa, 2025, Zhai et al., 2023, Gabrawy et al., 2022, Hermann et al., 2022, Gamez et al., 2021, Brody et al., 2020, Chipman, 2020, Koshikawa, 2020, Newell et al., 2020, Yusuff et al., 2020, Held and Sessions, 2019, Waghmare and Page-McCaw, 2018, Wiese et al., 2018, Anvarian et al., 2016, Chavez et al., 2016, Mottier-Pavie et al., 2016, Reid and O'Brochta, 2016, Urbach et al., 2016, Verghese and Su, 2016, Wieschaus and Nüsslein-Volhard, 2016, Auer et al., 2015, Bieli et al., 2015, Drusenheimer et al., 2015, Matsuda et al., 2015, Matsuda et al., 2015, Mishra et al., 2015, Pasco et al., 2015, Sugie et al., 2015, Verhulst and van de Zande, 2015, Verma and Cohen, 2015, Zhang et al., 2015, Alexandre et al., 2014, Herz et al., 2014, Ikmi et al., 2014, Kerr et al., 2014, Marr et al., 2014, Shukla et al., 2014, Singh and Mishra, 2014, Tower et al., 2014, Aoyama et al., 2013, Bejsovec, 2013, Das et al., 2013, Deshpande et al., 2013, Doumpas et al., 2013, Hombría and Sotillos, 2013, Howlett et al., 2013, Katsuyama et al., 2013, Konsavage and Yochum, 2013, Marques-Pita and Rocha, 2013, Ramos and Barolo, 2013, Stern and Frankel, 2013, Suzanne and Steller, 2013, Tan et al., 2013, Weasner and Kumar, 2013, Zoranovic et al., 2013, Ayukawa et al., 2012, Bier and Guichard, 2012, Formaz-Preston et al., 2012, Foronda et al., 2012, Gonsalvez and Long, 2012, Gurudatta et al., 2012, Hironaka et al., 2012, Jansen and Niessing, 2012, Kuroda et al., 2012, Pirooznia et al., 2012, Rincon-Limas et al., 2012, Swarup and Verheyen, 2012, White et al., 2012, Xie et al., 2012, Apidianakis and Rahme, 2011, Becam et al., 2011, Dalton et al., 2011, de Navas et al., 2011, Esteve et al., 2011, Gangaraju et al., 2011, Kryuchkov et al., 2011, Kurth et al., 2011, Ntini and Wimmer, 2011, O'Keefe et al., 2011, Repiso et al., 2011, Slattery et al., 2011, Stephan et al., 2011, Baig et al., 2010, Biehs et al., 2010, Braid et al., 2010, Cordero and Cagan, 2010, de Celis and Molnar, 2010, Gettings et al., 2010, Herranz et al., 2010, Jones et al., 2010, Kleinschmit et al., 2010, Kugler and Nagel, 2010, Lee et al., 2010, Liu et al., 2010, Mukai et al., 2010, Rendic et al., 2010, Roignant et al., 2010, Saj et al., 2010, Swaminathan et al., 2010, Wang et al., 2010, Werner et al., 2010, Williams et al., 2010, Yavari et al., 2010, Zecca and Struhl, 2010, Zheng et al., 2010, Baena-Lopez et al., 2009, Dichtel-Danjoy et al., 2009, Fang et al., 2009, Fre et al., 2009, Gaziova and Bhat, 2009, Gutierrez-Aviño et al., 2009, Martín et al., 2009, Martinez et al., 2009, Mulinari and Häcker, 2009, Nicholson et al., 2009, Ohayon et al., 2009, Perea et al., 2009, Roegiers et al., 2009, Widmann and Dahmann, 2009, Adan et al., 2008, Beckervordersandforth et al., 2008, Chen et al., 2008, dos Santos et al., 2008, Franch-Marro et al., 2008, Franch-Marro et al., 2008, Fromental-Ramain et al., 2008, Haussmann et al., 2008, Herranz et al., 2008, Hittinger and Carroll, 2008, Ishihara and Shibata, 2008, Ivanov et al., 2008, Larson et al., 2008, Leatherman and DiNardo, 2008, McClure et al., 2008, Peterson-Nedry et al., 2008, Sánchez et al., 2008, Shlevkov and Morata, 2008, Takashima et al., 2008, Ayala-Camargo et al., 2007, Ayala et al., 2007, Baig-Lewis et al., 2007, Bras-Pereira and Casares, 2007, Chao et al., 2007, DasGupta et al., 2007, Guan et al., 2007, Hatini and Nusinow, 2007, Hatton-Ellis et al., 2007, Kankel et al., 2007, Kondo et al., 2007, Lechner et al., 2007, Lee et al., 2007, Lindner et al., 2007, Loncle et al., 2007, Maeda et al., 2007, Najand and Simmonds, 2007, Nekrasov et al., 2007, Ruel et al., 2007, Samsonova et al., 2007, Tanaka et al., 2007, Tsai et al., 2007, Vendra et al., 2007, Vrailas-Mortimer et al., 2007, Wang and Gergen, 2007, Wang et al., 2007, Zirin and Mann, 2007, Zuckerkandl and Cavalli, 2007, Bowler et al., 2006, Childress et al., 2006, Cho et al., 2006, Deutsch, 2006, Eissenberg, 2006, Friedrich, 2006, Fuwa et al., 2006, Gallet et al., 2006, Hashimoto and Yamaguchi, 2006, Langdon et al., 2006, Martin and Morata, 2006, Matakatsu and Blair, 2006, Maurange et al., 2006, Parker, 2006, Pereira et al., 2006, Schwartz et al., 2006, Simpson et al., 2006, Tucker and Chiquet-Ehrismann, 2006, Wheeler et al., 2006, Wodarz et al., 2006, Ayyub et al., 2005, Davis et al., 2005, Hayward et al., 2005, Jordan et al., 2005, Martinez Arias, 2005, Pearson et al., 2005, Roederer et al., 2005, Takanaka and Courey, 2005, Linder and Deschenes, 2004, Trainor, 2004, Yang et al., 2004, Micchelli et al., 2003, Ludlam et al., 2002, Wilkie et al., 2001, Lawrence et al., 2000, Rogina and Helfand, 2000, Weinmaster, 1997, Sharma, 1973)
    Secondary FlyBase IDs
    • FBgn0004009
    • FBgn0001109
    • FBgn0003467
    • FBgn0003469
    • FBgn0011783
    • FBgn0065481
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    Study result (0)
    Result
    Result Type
    Title
    External Crossreferences and Linkouts ( 76 )
    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 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
    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
    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
    Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
    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
    iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
    Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
    KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    References (3,600)