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General Information
Symbol
Dmel\hh
Species
D. melanogaster
Name
hedgehog
Annotation Symbol
CG4637
Feature Type
FlyBase ID
FBgn0004644
Gene Model Status
Stock Availability
Gene Snapshot
hedgehog (hh) encodes the Hh signaling pathway ligand. It acts as a morphogen contributing to segment polarity determination, stem cells maintenance and cell migration. Post-translational modifications of the product of hh are essential for its restrictive spreading and signaling activity. [Date last reviewed: 2019-03-07]
Also Known As
Mrt, bar-3, l(3)neo56
Key Links
Genomic Location
Cytogenetic map
Sequence location
3R:23,128,169..23,141,906 [-]
Recombination map
3-78
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the hedgehog family. (Q02936)
Summaries
Pathway (FlyBase)
Hedgehog Signaling Pathway Core Components -
The hedgehog signaling pathway is initiated by hedgehog (hh) ligand binding to the extracellular domain of patched receptor (ptc), leading to the derepression of smoothened (smo) activity. Activation of the atypical GPCR smo results in the accumulation of the transcriptional activator form of cubitus interruptus (ci) and the derepression/activation of hh target genes. In the absence of hh, smo is repressed by ptc and ci is processed to a truncated repressor form. (Adapted from FBrf0220683 and FBrf0231236).
Gene Group (FlyBase)
UNCLASSIFIED RECEPTOR LIGANDS -
Receptor ligands are endogenous polypeptides that bind transmembrane cell surface receptors to elicit a cellular response.
Protein Function (UniProtKB)
Intercellular signal essential for a variety of patterning events during development. Establishes the anterior-posterior axis of the embryonic segments and patterns the larval imaginal disks. Binds to the patched (ptc) receptor, which functions in association with smoothened (smo), to activate the transcription of target genes wingless (wg), decapentaplegic (dpp) and ptc. In the absence of hh, ptc represses the constitutive signaling activity of smo through fused (fu). Essential component of a signaling pathway which regulates the Duox-dependent gut immune response to bacterial uracil; required to activate Cad99C-dependent endosome formation, norpA-dependent Ca2+ mobilization and p38 MAPK, which are essential steps in the Duox-dependent production of reactive oxygen species (ROS) in response to intestinal bacterial infection (PubMed:25639794). During photoreceptor differentiation, it up-regulates transcription of Ubr3, which in turn promotes the hh-signaling pathway by mediating the ubiquitination and degradation of cos (PubMed:27195754). The hedgehog protein N-product constitutes the active species in both local and long-range signaling, whereas the C-terminal product has no signaling activity. It acts as a morphogen, and diffuses long distances despite its lipidation. Heparan sulfate proteoglycans of the extracellular matrix play an essential role in diffusion. Lipophorin is required for diffusion, probably by acting as vehicle for its movement, explaining how it can spread over long distances despite its lipidation. The hedgehog protein C-product, which mediates the autocatalytic activity, has no signaling activity.
(UniProt, Q02936)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
hh: hedgehog
A segment polarity type of embryonic lethal. Homozygous embryos have the posterior naked portion of the ventral surface of each segment deleted and replaced by a mirror image of the anterior denticle belts. Embryos appear to lack segmental boundaries. In strong alleles, there is no obvious segmentation; the larvae are approximately 40% the length of the wild-type larvae, and there is a lawn of denticles arranged in a number of whorls on the ventral surface as a result of loss of naked cuticle. In intermediate alleles, naked cuticle is also lost from the ventral region, but the lawn of denticles is arranged in segmental arrays in mirror-image symmetry. The weak alleles show fusions that delete the naked cuticle usually between abdominal segments 1 and 2 and 6, 7, and 8 (Mohler, 1988). Temperature shift experiments with a temperature-sensitive allele (viable and normal at 18, and mutant at 25) indicate two phases of hh activity at 25, the first during early embryogenesis (3-6 hr of development) and the second during the late larval and early pupal stages (4-7 days of development).
hh1
A weak hypomorphic allele that is not complemented by other hh alleles. Eye of homozygote small and narrow with about 150 facets. Eye disc size reduced; deep cleft at anterior edge cell; clusters at cleft look mature (Renfranz and Benzer, 1989, Dev. Biol. 136: 411-29).
Mir: Mirabile (M. Muskavitch)
Mirror-image duplication of tergite structure. Microchaetae are eliminated from the anterior portion of the tergite and replaced by a duplication consisting of an anteriorly oriented row of macrochaetae and the darkly pigmented cuticle normally found in the posterior portion of the tergite. Fat body and oenocytes underneath the tergite are also duplicated with mirror-image symmetry (Madhavan and Madhavan).
Mrt: Moonrat (J.A. Kennison)
Heterozygote shows partial transformation of anterior wing to posterior (triple row bristles replaced by double row bristles in patches). A network of extra veins appears in the anterior compartment, beginning at the distal edge in the least affected flies, and covering the entire anterior compartment in the more extreme cases. Wing blade expanded anteriorly at the distal edge. Wing blade expansion and extra veins resemble phenotypes seen in en1 homozygotes in the presence of Minute mutations. Bubbles often form in the wing blade. More rarely, a mirror-image outgrowth from the anterior edge is present. Mirror-image duplications sometimes appear in halteres. Legs sometimes appear deformed (similar to phenotype of enlethal clones induced in the larva). Dominant phenotypes strongly temperature-sensitive. Penetrance greater than 99% at 18 (with strong expressivity) but only 30-40% at 29 (with very weak expressivity). Shows paternal effect. Penetrance greater when mutant allele inherited from father than when inherited from mother. Mrt/+/+ indistinguishable from Mrt/+.
Summary (Interactive Fly)
Hedgehog N-terminal signaling domain and C-terminal autoprocessing domain - helps establish embryonic segmentation - a segment polarity intercellular signaling protein - cooperates with Frazzled to guide axons through a non-canonical signalling pathway - A local difference in Hedgehog signal transduction increases mechanical cell bond tension and biases cell intercalations along the Drosophila anteroposterior compartment boundary
Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\hh or the JBrowse view of Dmel\hh 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
Comments on Gene Model
Gene model reviewed during 5.44
Gene model reviewed during 5.48
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0100506
2370
471
Additional Transcript Data and Comments
Reported size (kB)
2.3 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0099945
52.1
471
8.23
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)
471 (aa); 52 (kD)
471 (aa); 52 (kD predicted)
Comments
External Data
Subunit Structure (UniProtKB)
Interacts with shf.
(UniProt, Q02936)
Post Translational Modification
The C-terminal domain displays autoproteolytic activity. Cleavage of the full-length hedgehog protein is followed by the covalent attachment of a cholesterol moiety to the C-terminus of the newly generated N-terminal fragment (N-product). Cholesterol attachment plays an essential role in restricting the spatial distribution of hedgehog activity to the cell surface. N-terminal palmitoylation of the hedgehog N-product is required for the embryonic and larval patterning activities of the hedgehog signal. Rasp acts within the secretory pathway to catalyze the N-terminal palmitoylation of Hh.
(UniProt, Q02936)
Crossreferences
MEROPS - An information resource for peptidases (also termed proteases, proteinases and proteolytic enzymes) and the proteins that inhibit them.
PDB - An information portal to biological macromolecular structures
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

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

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (73 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:ihog; FB:FBgn0031872
inferred from physical interaction with UniProtKB:Q9VM64
(assigned by UniProt )
inferred from physical interaction with UniProtKB:Q9W3W5
(assigned by UniProt )
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN001726121
(assigned by GO_Central )
non-traceable author statement
(assigned by UniProt )
traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN001726121
(assigned by GO_Central )
inferred from electronic annotation with InterPro:IPR001767
(assigned by InterPro )
non-traceable author statement
(assigned by UniProt )
Biological Process (59 terms)
Terms Based on Experimental Evidence (33 terms)
CV Term
Evidence
References
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from expression pattern
(assigned by UniProt )
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
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 direct assay
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:disp; FB:FBgn0029088
inferred from genetic interaction with FLYBASE:Hmgcr; FB:FBgn0263782
inferred from expression pattern
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:dpp; FB:FBgn0000490
inferred from genetic interaction with FLYBASE:h; FB:FBgn0001168
inferred from genetic interaction with FLYBASE:sca; FB:FBgn0003326
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
inferred from mutant phenotype
Terms Based on Predictions or Assertions (31 terms)
CV Term
Evidence
References
traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN001726121
(assigned by GO_Central )
non-traceable author statement
traceable author statement
traceable author statement
traceable author statement
traceable author statement
traceable author statement
traceable author statement
inferred from electronic annotation with InterPro:IPR006141
(assigned by InterPro )
non-traceable author statement
traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN001726121
(assigned by GO_Central )
traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN001726121
(assigned by GO_Central )
traceable author statement
Cellular Component (8 terms)
Terms Based on Experimental Evidence (8 terms)
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
inferred from direct assay
inferred from direct assay
inferred from direct assay
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
non-traceable author statement
(assigned by UniProt )
traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN001726121
(assigned by GO_Central )
non-traceable author statement
(assigned by UniProt )
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
hh RNA can be detected in ganglion mother cells at 72 hours after larval hatching (ALH), but not at 48 hours ALH.
hh is expressed strongly in the anterior escort cells, and weakly in the posterior escort cells
At early stage 11, hh expression extends from the posterior optic lobe into the anterior optic lobe. It becomes restricted to the posterior optic lobe at mid stage 11 and later localizes to Bolwig organ precursor cells in the ventral posterior optic lobe. hh is expressed in cells immediately anterior to the tracheal pits.
hh transcripts are expressed in embryos, larvae, pupae and adults with peaks of expression in 6-12hr embryos and early pupae. Transcripts are first detected in embryonic stage 5 in a few stripes at the anterior and posterior ends of the embryo. The number of stripes gradually increases to 17. The terminal stripes are 2-3 cells wide and the internal stripes are 1 cell wide. Expression is stronger in every second stripe and is stronger laterally than in dorsal and ventral regions. The expression in stripes reaches a maximum at stages 8-11. By the end of germband retraction, the stripes are situated in the posterior compartments of the lateral ectoderm.
Peaks of hh expression are observedin 2-10hr embryos and in pupae. In embryos, hh transcripts are firstexpressed in a discrete pattern in the maxillary segment followed by apattern of 14 parasegmental stripes. At germ band extension, a 15th stripeis seen. hh expression in the metameric portion of the embryo closelyresembles en expression. Expression is also described in a variety ofsites in the nonmetameric portion of the embryo including the intercalaryand antennal segments, the procephalon, the gnathal segments, and portionsof the hindgut. Expression in imaginal discs is described for theassociated Ecol\lacZ insertion. hh expression is pair-rule dependent. Inftz mutants, expression in the even-numbered parasegments is missing.wg mutations caused diminished expression and ptc mutants causeexpression in an ectopic stripe in each segment. nkd mutations causebroadening of the stripes.
hh transcripts are first detected at the cellular blastoderm stage in 17 segmental stripes. 14 of the stripes are one cell wide and extend from 10-70% egg length. There are two 3-cell-wide stripes at 5% and 75% egg length and a dorsal anterior spot at 97% egg length. The stripes appear asynchronously. Even parasegmentally-numbered stripes precede odd-numbered stripes and anterior stripes precede more posterior stripes. The stripes are activated around the entire circumference of the embryo but disappear from the amnioserosa and mesoderm after gastrulation. At stage 11, the stripes are located just posterior to the parasegmental furrow and are spaced one cell anterior to the tracheal pit in each segment. Stripes persist after germ band retraction and are located in the posteriormost portion of the lateral ectoderm of each segment. hh transcripts are also expressed in the fore- and hindguts following gastrulation and germ band extension as well as in the cephalic region of the embryo. Up to stage 10, the intensity of staining is heavier in the nucleus than in the cytoplasm. After stage 10, RNA staining is predominantly cytoplasmic.
At the cellular blastoderm stage, hh transcripts are located predominantly in a single stripe at 75% egg length with additional transcripts at the anterior tip and along the ventral side. At gastrulation, hh is expressed in 14 single-cell-wide stripes between 30% and 65% egg length. The stripes are coincident with en expression and occur in the cells of the posterior compartments. They appear in a characteristic order, even-numbered ones before odd-numbered ones and anterior ones before posterior ones. hh is also expressed in a block of cells at the anterior end and in wide stripes at 10% and 75% egg length. hh transcripts are expressed later in the foregut, pharynx, esophagus, hindgut, and salivary glands. hh transcripts are exressed in imaginal discs where they are localized to the posterior compartments. The differences between hh and en expression are noted.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
hh protein is expressed at similar levels in wing and haltere discs.
Expression of hh in the eye-antennal disc is more restricted than the expression of ptc. hh protein is present in cells adjacent to ptc-expressing cells in only certain regions.
Marker for
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
inferred from direct assay
inferred from direct assay
inferred from direct assay
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
Expression Deduced from Reporters
Reporter: P{GAL4}hhGal4
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lacW}A937A
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lwB}16E
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lwB}hhH90
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{PZ}hhP30
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{PZ}hhQ50
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{PZ}hhrJ413
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{PZ}P2023-44
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\hh in GBrowse 2
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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, and Transgenic Constructs
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
abdominal tergite & macrochaeta | conditional ts
abdominal tergite & microchaeta
abdominal tergite & microchaeta | conditional ts
abdominal tergite | anterior & macrochaeta
abdominal tergite | anterior & microchaeta
abdominal tergite | anterior & trichome
adult cuticle & head capsule | dorsal | conditional ts
cytoneme & dorsal mesothoracic disc | somatic clone
embryonic abdominal segment 1 & cuticle, with Scer\GAL4prd.RG1
embryonic abdominal segment 1 & denticle belt, with Scer\GAL4prd.RG1
embryonic abdominal segment 3 & denticle belt, with Scer\GAL4prd.RG1
embryonic abdominal segment 3 & denticle belt | supernumerary, with Scer\GAL4prd.RG1
embryonic abdominal segment 5 & denticle belt, with Scer\GAL4prd.RG1
embryonic abdominal segment 5 & denticle belt | supernumerary, with Scer\GAL4prd.RG1
embryonic abdominal segment 7 & denticle belt, with Scer\GAL4prd.RG1
embryonic abdominal segment 7 & denticle belt | supernumerary, with Scer\GAL4prd.RG1
embryonic thoracic segment & cuticle, with Scer\GAL4prd.RG1
embryonic thoracic segment & denticle belt, with Scer\GAL4prd.RG1
eye (with hh8)
eye (with hhbar3)
eye & ommatidium
glial cell & eye disc | somatic clone | cell non-autonomous, with Scer\GAL4Act5C.PP
lamina & neuron
lamina & neuron | precursor
macrochaeta & tarsal segment 5 | distal
microchaeta & tarsal segment 1
microchaeta & tarsal segment 2
microchaeta & tarsal segment 3
microchaeta & tarsal segment 4
microchaeta & tarsal segment 5
microchaeta & wing | anterior | proximal
oocyte & microtubule
photoreceptor cell & axon
photoreceptor cell R7 & axon
photoreceptor cell R8 & axon
scutellum & macrochaeta
scutellum & macrochaeta, with Scer\GAL4C-734
scutellum & macrochaeta, with Scer\GAL4en-e16E
wing (with hhMrt)
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (3)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
14 of 15
Yes
Yes
13 of 15
No
Yes
 
12 of 15
No
Yes
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (3)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
13 of 15
Yes
Yes
13 of 15
Yes
Yes
 
11 of 15
No
Yes
Rattus norvegicus (Norway rat) (3)
12 of 13
Yes
Yes
9 of 13
No
Yes
7 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (3)
8 of 12
Yes
Yes
6 of 12
No
Yes
6 of 12
No
Yes
Danio rerio (Zebrafish) (5)
13 of 15
Yes
Yes
12 of 15
No
Yes
11 of 15
No
Yes
9 of 15
No
Yes
6 of 15
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (15)
5 of 15
Yes
Yes
4 of 15
No
Yes
4 of 15
No
Yes
4 of 15
No
Yes
4 of 15
No
Yes
4 of 15
No
Yes
4 of 15
No
Yes
4 of 15
No
Yes
3 of 15
No
Yes
2 of 15
No
Yes
2 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
Yes
Arabidopsis thaliana (thale-cress) (1)
1 of 9
Yes
Yes
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091909F1 )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG09150573 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0EJW )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0EJ3 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0N90 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (0)
No records found.
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Allele
Disease
Evidence
References
Potential Models Based on Orthology ( 4 )
Modifiers Based on Experimental Evidence ( 4 )
Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
 
Disease Associations of Human Orthologs (via DIOPT v7.1 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
esyN Network Diagram
Show neighbor-neighbor interactions:
Select Layout:
Legend:
Protein
RNA
Selected Interactor(s)
Interactions Browser

Please see the Physical Interaction reports below for full details
protein-protein
Physical Interaction
Assay
References
Summary of Genetic Interactions
esyN Network Diagram
esyN Network Key:
Suppression
Enhancement

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
enhanceable
suppressible
suppressible
External Data
Subunit Structure (UniProtKB)
Interacts with shf.
(UniProt, Q02936 )
Linkouts
BioGRID - A database of protein and genetic interactions.
DroID - A comprehensive database of gene and protein interactions.
InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Gene Group - Pathway Membership (FlyBase)
Hedgehog Signaling Pathway Core Components -
The hedgehog signaling pathway is initiated by hedgehog (hh) ligand binding to the extracellular domain of patched receptor (ptc), leading to the derepression of smoothened (smo) activity. Activation of the atypical GPCR smo results in the accumulation of the transcriptional activator form of cubitus interruptus (ci) and the derepression/activation of hh target genes. In the absence of hh, smo is repressed by ptc and ci is processed to a truncated repressor form. (Adapted from FBrf0220683 and FBrf0231236).
External Data
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3R
Recombination map
3-78
Cytogenetic map
Sequence location
3R:23,128,169..23,141,906 [-]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
94E1-94E1
Limits computationally determined from genome sequence between P{lacW}GclmL0580 and P{EP}hhEP3521
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
94E1-94E3
(determined by in situ hybridisation)
94E2-94E2
(determined by in situ hybridisation)
94E1-94E4
(determined by in situ hybridisation)
94D-94E
(determined by in situ hybridisation)
94D10-94E5
(determined by in situ hybridisation)
94D10-94D13
(determined by in situ hybridisation)
On the basis of meiotic mapping.
Experimentally Determined Recombination Data
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (38)
Genomic Clones (13)
 

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

cDNA Clones (4)
 

Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see GBrowse for alignment of the cDNAs and ESTs to the gene model.

cDNA clones, fully sequences
BDGP DGC clones
    Other clones
      Drosophila Genomics Resource Center cDNA clones

      For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

      cDNA Clones, End Sequenced (ESTs)
      BDGP DGC clones
        Other clones
          RNAi and Array Information
          Linkouts
          DRSC - Results frm RNAi screens
          GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
          Antibody Information
          Laboratory Generated Antibodies
          Commercially Available Antibodies
           
          Other Information
          Relationship to Other Genes
          Source for database identify of
          Source for database merge of
          Source for merge of: hh l(3)neo56
          Source for merge of: hh anon-WO0134654.19
          Additional comments
          Source for merge of hh anon-WO0134654.19 was sequence comparison ( date:051113 ).
          Other Comments
          Cholesterol modification of hh is necessary for hh-dependent graded cell fate specification in the dorsal epidermis.
          One of 42 Drosophila genes identified as being most likely to reveal molecular and cellular mechanisms of nervous system development or plasticity relevant to human Mental Retardation disorders.
          Cholesterol modification of hh protein is necessary (in the embryonic epidermis) for its assembly in large punctate subcellular structures and apical sorting through the activity of the disp protein. Movement of these specialized structures containing hh protein through the cellular field is contingent upon the activity of ttv.
          hh is responsible for maintaining Con expression and its own expression in the embryonic trunk mesoderm during gut and tracheal development.
          gish, so, ey and hh act in the posterior region of the eye disc to prevent precocious glial cell migration.
          The expression of hh in the wing disc, once activated, is dependent on ph-p, trx and brm. This may be due to an element upstream of the hh transcriptional start site (hh-CMM) that can bind Pc protein and is able to act as a cellular memory module (CMM) when placed upstream of a UAS sequence in reporter constructs.
          Misexpression of hh in the soma induces germ cells to migrate to inappropriate locations in the developing embryo.
          hh may act as an attractive guidance cue in germ cell migration in the embryo.
          Migration of all tracheal branches is absent or stalled in hh-mutant embryos.
          hh expression in the prospective rectum is necessary for the expression of dpp at the posterior end of the adjacent large intestine. hh expression is also required for the development of the rectum.
          hh acts as a somatic stem cell factor in the ovary.
          Clones of hh mutants in the peripodial membrane disrupt disc growth.
          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.
          hh signalling from the peripodial membrane, but not from the disc proper, is required for eye disc patterning and growth.
          ptc protein destabilises smo protein in the absence of hh protein.
          Bolwig's organ formation is governed by ato, the expression of which is under the control of hh, eya and so.
          hh activates vn expression. This activation is mediated through the gene ci.
          hh induces Egfr signalling during head development.
          hh is required in the developing eye both for the induction of ato expression that prefigures the position of the R8 cells, and for the repression of ato expression between the nascent proneural clusters. Both effects are due to the direct stimulation of responding cells by the hh gene product itself.
          hh plays a role in ommatidial development by regulating ato expression both positively and negatively.
          In the absence of hh activity, prothoracic leg disc fragments fail to undergo anterior/posterior conversion, but can still regenerate missing anterior pattern elements. hh-independent regeneration (integration) may be mediated by the wg and dpp positional cues.
          hh is required for activation of en during regeneration of fragmented imaginal discs.
          The en/hh interface in the embryonic epidermis imposes asymmetry on wg signaling.
          Three EMS induced alleles were identified in a screen for mutations affecting commissure formation in the CNS of the embryo.
          hh signalling, coming from the adjacent P compartments across both Anterior/Posterior and Posterior/Anterior boundaries in the abdomen, organizes the pattern of all the Anterior cells.
          Cell affinities in the adult abdomen depend on hh : cells of the A compartment show two gradients of affinity, both of which depend on direct readouts of the level of hh function.
          cad acts in combination with the hh pathway to specify the different components of the analia.
          hh plays a role in the formation of the posterior barrier to wg movement at the presumptive embryonic segment border. Increased hh signalling decreases the domain of wg action in the anterior direction.
          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 hh signalling components smo and ci are required in cells posterior to hh to maintain ptc expression, whereas fu is not necessary in these cells.
          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.
          hh is required at the posterior margin of the eye disc to maintain expression of dpp and ato.
          ptc protein normally binds hh gene product without any help of the smo gene product, though smo is also a part of the receptor complex that binds hh and transduces the hh signal. The mechanism of signal transduction may involve hh binding specifically to ptc and inducing a conformational change leading to the release of latent smo activity.
          The division of the limb into two antagonistic domains, as defined by exd function and hh signaling, may be a general feature of limb development.
          hh and spi bring about the concerted assembly of ommatidial and synaptic cartridge units, imposing the "neurocrystalline" order of the compound eye onto the post-synaptic target field. hh encodes an inductive signal that is transported along retinal axons from the developing eye, and induces the expression of Egfr in post-synaptic precursor cells.
          In the absence of hh signalling results propose that Su(fu) inhibits ci by binding to it and that, upon reception of the hh signal, fu is activated and counteracts Su(fu), leading to the activation of ci.
          Mutants are isolated in an EMS mutagenesis screen to identify zygotic mutations affecting germ cell migration at discrete points during embryogenesis: mutants exhibit segment polarity pattern defects.
          The hh product stimulates maturation of ci into a labile transcriptional activator.
          CrebA hh double mutant phenotype confirms that CrebA is not involved in segment polarity.
          Processing of the full length ci protein is inhibited by hh, an observation that represents the first direct evidence that ci transduces the hh signal.
          Each primordia of the genital disc (female genital, male genital and anal primordia) is divided into anterior and posterior compartments. 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.
          Genetic combinations with mutants of nub cause additive phenotypes.
          Clonal analysis demonstrates hh has two distinct functions: expression is required in the photoreceptor cells to drive the morphogenetic furrow and in addition hh secreted from cells at the posterior disc margin is absolutely required for the initiation of patterning and predisposes ommatidial precursor cells to enter ommatidial assembly later.
          hh induces ommatidial development in the absence of its secondary signals wg and dpp. Regulatory relationships between hh, dpp and wg in the eye are similar to those found in other imaginal discs, such as the leg.
          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.
          Identified in a screen for modifiers of the Dfd13/Dfd3 mutant phenotype. Shows no interaction with the Pc mutant phenotype.
          ci forms a negative feedback loop with ptc that is regulatd by hh signal transduction.
          bi is the primary target of hh signaling in the adult abdomen, mediating both the morphogenetic and polarity-reversal functions of hh.
          hh protein secreted by posterior compartment cells plays a key role in patterning the posterior portion of the anterior compartment in adult abdominal segments.
          dpp specifies the position of most of the sensory organ precursors (SOPs) in the notum and some of them in the wing. Close to the A/P compartment border of the wing, however, SOPs are specified by hh rather than by dpp alone.
          dpp only mediates a subset of hh functions in the morphogenetic furrow.
          dpp does not appear to be the principal mediator of hh function in the eye.
          Loss of smo function causes a hh-like phenotype. smo activity is required for transduction of hh but not wg. smo acts downstream from ptc to transduce the hh signal.
          hh elicits signal transduction via a complex that includes the products of the fu, ci and cos genes. The complex binds with high affinity to microtubules in the absence of hh protein, but not when hh is present. The complex may facilitate signalling from hh by governing access of the ci product to the nucleus.
          The affinity boundary that segregates A and P cells into adjacent but immiscible cell populations is to a large extent a consequence of local hh signalling, rather than a reflection of an intrinsic affinity difference between A and P cells.
          cos encodes a kinesin-related protein that accumulates preferentially in cells capable of responding to hh signal.
          Comparing the biological activities of secreted and membrane-tethered hh protein provides evidence that hh forms a local concentration gradient and functions as a concentration-dependent morphogen in the wing.
          The pattern of expression of hh in the larval and adult abdomen has been analysed.
          The function of hh in morphogenetic furrow progression is indirect. Cells that cannot receive/transduce the hh signal (as in smo clones) are still capable of entering a furrow fate and differentiating normally. However hh is required to promote furrow progression and regulate its rate of movement across the disc, since the furrow is delayed in smo clones.
          hh and ptc can regulate transcription from a wg enhancer element containing ci protein binding sites by modulating the activity of ci protein.
          smo encodes a seven-pass membrane protein, a putative receptor of the hh signal.
          Elevated levels of ci are sufficient to activate hh target genes, even in the absence of hh activity. ci activates transcription in yeast by a GLI consensus-binding site and the zinc finger domain is sufficient for its target specificity. Results strongly support a role for ci as the transcriptional activator that mediates hh signaling.
          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. hh and wg play opposing roles in mesoderm segmentation.
          wg and hh signaling account for all cell types across the dorsal epidermis. dpp does not appear to mediate this hh dorsal epidermis signaling. hh antagonizes the activity of ptc in the specification of primary and secondary but not tertiary cell types. hh also antagonizes lin function.
          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.
          Loss of da disrupts the progression of the morphogenetic furrow and this effect is mediated by the loss of both hh and dpp.
          smo activity is required in wing anterior cells along the A/P boundary for these cells both to transduce hh and to limit its further movement into the anterior compartment. ptc regulates smo activity in response to hh signalling.
          Ectodermal and mesodermal Dr expression depend on wg and hh.
          Cells in anterior compartments lacking ci express hh and adopt a posterior fate without expressing en. Increased levels of ci can induce the expression of dpp independent of hh. Expression of ci in anterior cells controls limb development by restricting hh secretion to posterior cells and by conferring competence to respond to hh by mediating transduction of the hh signal.
          ptc and ci are expressed in a pattern complementary to hh and en in adult ovaries. Ectopic expression of hh results in the ectopic expression of ptc. hh directly effects region 2 somatic cells of the germarium via a signalling pathway which includes ptc and ci, but not wg or dpp.
          hh is required for the proliferation and specification of ovarian somatic cells prior to egg chamber formation. hh signalling during egg chamber assembly appears to be closley related to, or part of pathways involving the neurogenic genes.
          The expression pattern of a number of genes in the larval genital discs, including a hh-Ecol\lacZ reporter gene, has been studied to determine the segment-parasegment organisation of the genital discs.
          ara-caup expression at patches on the wing, located one at each side of the DV compartment border, is mediated by the hh signal through its induction of high levels of ci protein in anterior cells near to the AP compartment border.
          exd is expressed in a normal pattern in the absence of hh function.
          hh, wg and dpp are required for the establishment of signaling centres that coordinate morphogenesis in the hindgut epithelium. Activation of these genes 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.
          The secreted hh product regulates the temporal assembly of photoreceptor precursor cells into ommatidia in the eye and is transmitted along the retinal axons to serve as the inductive signal in the brain, triggering neurogenesis in the developing visual centers. hh acts in the first of two retinal axon-mediated steps in the assembly of lamina synaptic cartridges.
          A combination of hh and wg is required to specify the most posterior fates of the A compartment.
          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).
          hh is required in the early gastrula for heart development, overexpression of hh increases the amount of heart formation. Overexpression of wg restores the heart deficit of hh mutant embryos.
          The hh autoprocessing reaction proceeds via an internal thioester intermediate and results in a covalent modification that increases the hydrophobic character of the signalling domain and influences its spatial and subcellular distribution. Truncated, unprocessed amino terminal protein causes embryonic mispatterning, suggesting a role for autoprocessing in spatial regulation of hh signalling.
          Cholesterol is the lipophilic moiety covalently attached to the amino-terminal signalling domain during autoprocessing. The carboxy-terminal domain acts as an intramolecular cholesterol transferase.
          hh and wg specify the identities of specific regions of the head capsule. 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. oc is a candidate hh target gene during early eye-antennal disc development.
          In competition binding, cross-linking and co-immunoprecipitation experiments no binding of tagged hh protein to smo protein or its rat homolog could be detected, although hh protein can bind to the protein encoded by the mouse homolog of ptc.
          en is not required for hh activation or maintenance in the eye imaginal disc.
          fu protein is phosphorylated during embryogenesis as a result of hh activity. Results from cell culture studies suggest that fu and Pka-C1 function downstream of hh but in parallel pathways that eventually converge distal to fu.
          Segment polarity gene smo is required for the response of cells to hh signalling during the development of both the embryonic segments and imaginal discs. Structure of the smo protein suggests it may act as a receptor for the hh ligand.
          The small lobe of fu may play a role in generating the neomorphic cos phenotype displayed by an unregulated fu protein in a Su(fu)- background.
          hh protein acts in the wing as a signal to instruct neighbouring cells to adopt fates appropriate to the region of the wing just anterior to the compartmental boundary. Some members of the trx group genes are involved in the transcriptional regulation of genes in the hh signalling pathway during imaginal development, Pc group genes are not involved in this regulation pathway.
          Ectopic expression of the amino-terminal half of the hh protein results in effects similar to those induced by the wild-type protein, altering the identity of cells of both the dorsal and ventral ectoderm of the developing embryo and of cells of the anterior compartment of the imaginal discs. Ectopic expression of a form of the protein in which the signal cleavage sequence is mutated has no effect on larval or adult development. Results suggest that all signaling activity of the hh protein is most likely to reside in the amino terminal fragment generated by autoproteolysis.
          Mutations of hh interact with Dfd to reduce the viability of the Dfd3/Dfd13 combination.
          Ectopic expression of hh produces ectopic furrows in the anterior eye disc. In addition to changes in cell shape the ectopic furrows are associated cell proliferation, cell cycle synchronisation and pattern formation, events that parallel normal furrow progression. Results propose that the morphogenetic furrow coincides with a transient boundary that coordinates growth and differentiation of the eye disc and hh is necessary and sufficient to propagate this boundary across the epithelium.
          Ectopic hh causes respecification of the wing anterior compartment. Reorganisation of the anterior wing is presaged by ectopic expression of dpp and ptc.
          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.
          Pka-C1 activity is not regulated by ptc but may be regulated by hh.
          The effects of mutations in the anterior gap genes hkb, tll, oc, ems and btd on the spatial expression of hh and wg during embryogenesis have been investigated.
          fu and hh modulate the post-transcriptional regulation of ci protein.
          Pka-C1 is a component of the signalling pathway that represses dpp expression in the anterior compartment in appendage imaginal discs and anterior to the morphogenetic furrow in eye discs.
          hh, wg and mys are required for epithelial morphogenesis during proventriculus organ development. The morphogenetic process is suppressed by dpp. These results identify a novel cell signalling centre in the foregut that operates through a distinct genetic circuitry in the midgut to direct the formation of a multiply folded organ from a simple epithelial tube.
          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.
          hh pathway mutants induce ectopic morphogenetic furrows. Results show that ommatidial clusters are self-organising units whose polarity in one axis is determined by the direction of furrow progression and which can independently define the position of an equator without reference to the global coordinates of the eye disc.
          Both ptc and Pka-C1 act downstream of hh in the developing eye.
          en governs growth and patterning in both anterior and posterior wing compartments by controlling the expression of the hh and dpp products as well as the response of the cells to them. en activity programs wing cells to express hh whereas the absence of en activity programs them to respond to hh by expressing dpp. Consequently, posterior cells secrete hh product and induce a stripe of neighboring anterior cells across the compartment boundary to secrete dpp. dpp may exert its organizing influence by acting as a gradient morphogen in contrast to hh which appears to act principally as a short range inducer of dpp.
          gro and hh regulate en expression in the anterior compartment of the wing.
          Ectopic expression of hh can induce ectopic wg and dpp expression in anterior cells and reorganise the anterior compartment pattern. Loss of endogenous hh blocks wg and dpp expression along the compartment boundary and impedes growth and patterning in both compartments.
          Ectopic expression of hh in the anterior compartment of the wing disc causes overgrowth and pattern duplications in both anterior and posterior compartments of the wing disc, similar to alterations seen with ectopic dpp expression. These results indicate that hh is acting as a regulator of dpp expression and dpp acts as an organising molecule controlling growth and patterning in the wing imaginal disc.
          The maintenance of wg expression by the hh signal is limited to early development. In a later role, hh organizes segmental pattern acting in a concentration-dependent manner, as a morphogen.
          hh mediates the interaction between anterior dpp-expressing cells and posterior en-expressing cells.
          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.
          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.
          hh gene product is involved in regulating ptc expression in both embryo and discs, through its role in regulating gene expression along the anterior-posterior compartment border. hh function establishes the proximodistal axis in discs. hh protein is secreted and can cross embryo parasegment borders and the anterior-posterior compartment border of imaginal discs to neighbouring cells that express neither en nor hh. In the embryo hh regulation of ptc apparently facilitates ptc and wg expression. In the discs hh regulation of ptc and other genes in the anterior compartment helps to establish the proximodistal axis. Cell-cell communication mediated by hh links the special properties of compartment borders with specification of the proximodistal axis in imaginal development.
          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 and en is required for the first row of denticles in each abdominal denticle belt. Remaining cell types are produced by different combinations of the five gene activities. 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. hh and en contribute to the pattern independently. hh and ptc show mutual suppression through opposing effects on wg expression. hh alters the competence of cells to respond to wg signal.
          Transcriptional control of both ptc and wg by hh is mediated by the same signal transduction pathway.
          Developing retinal cells drive the progression of morphogenesis using the products of the hh and dpp genes. Clonal analysis suggests that gene products act as diffusible signals. hh induces the expression of dpp, the primary mediator of furrow movement.
          Segment polarity mutations cause stripes of abnormal patterning within sectors of the leg disc, which may be mediated by regional perturbations in growth.
          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. Maintenance of wg expression depends upon the activity of hh, which acts only on neighboring cells to maintain wg expression. Expression of wg in the absence of ptc depends on hh.
          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.
          The role of hh in the regulation of run mRNA expression in the early embryo has been investigated.
          A hh-related gene family has been identified in the zebrafish. Over-expression of one member, sonic hedgehog, in fly embryos, can activate the hh-dependent pathway.
          Many alleles of hh act as dominant alleles of gl.
          hh expression posterior to the morphogenetic furrow in the developing eye disc is continuously required for its progression. The forward diffusion of hh protein induces anterior cells to enter the furrow. hh acts upstream of gl, sca, h and dpp in the developing eye.
          Although hh is essential for wg function in segmentation, wg appears to be still capable of some action in hh's absence.
          Probably encodes a secreted or transmembrane protein.
          The pattern of hh protein expression during embryonic development has been analysed.
          wg and en expression patterns are studied in all known segment polarity mutants to investigate the requirement of other segment polarity genes in mediating the maintenance of wg and en.
          hh has been isolated and characterised.
          hh gene cloned by plasmid rescue, the encoded protein is targetted to the secretory pathway (consistant with the non-cell autonomous requirement for hedgehog in cuticular patterning) and is expressed coincidentally with engrailed in embryos and imaginal discs. Maintainance of hh expression is dependent upon other segment polarity genes including engrailed and wingless. The amino acid sequence shows no similarity to any known protein but hydropathy analysis highlights a prominent hydrophobic region.
          Sequence analysis of hh indicates that the gene product contains a putative transmembrane domain which suggests that it may be localized at the cell surface and be involved in cell-cell communication.
          hh gene cloned and the sequence suggests a membrane associated protein. Expression pattern analysed and found to coincide with that of en in the epidermis. Though initially independent of en, hh expression later becomes en-dependent.
          hh 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. Results suggest that ptc protein may be the receptor for the hh signal, implying that this is an usual mechanism of ligand-dependent receptor inactivation.
          hh is essential for maintaining the normal pattern of ptc expression.
          Role of hh in neurogenesis has been studied.
          Adult eyes are small, narrow with about 150 facets, eye disc is small due to precursor cell defects.
          The mutational effects of hh on larval and adult cuticular patterns has been studied. Defects in the distal portions of the legs and antenna occur in association with homozygous hh clones in the posterior compartments of the structures.
          Genetic mosaics were used to determine that hh is not autonomous at the level of the single cell.
          hh mutants display segment polarity segmentation defects.
          A segment polarity type of embryonic lethal. Homozygous embryos have the posterior naked portion of the ventral surface of each segment deleted and replaced by a mirror image of the anterior denticle belts. Embryos appear to lack segmental boundaries. In strong alleles, there is no obvious segmentation; the larvae are approximately 40% the length of the wild-type larvae and there is a lawn of denticles arranged in a number of whorls on the ventral surface as a result of loss of naked cuticle. In intermediate alleles, naked cuticle is also lost from the ventral region, but the lawn of denticles is arranged in segmental arrays in mirror-image symmetry. The weak alleles show fusions that delete the naked cuticle usually between abdominal segments 1 and 2 and 6, 7 and 8 (Mohler, 1988). Temperature shift experiments with a temperature-sensitive allele (viable and normal at 18oC, and mutant at 25oC) indicate two phases of hh activity at 25oC, the first during early embryogenesis (3-6 hr of development) and the second during the late larval and early pupal stages (4-7 days of development).
          Origin and Etymology
          Discoverer
          Etymology
          Identification
          External Crossreferences and Linkouts ( 70 )
          Sequence Crossreferences
          NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
          GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
          GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
          RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
          UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
          Other crossreferences
          Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
          Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
          Flygut - An atlas of the Drosophila adult midgut
          Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
          GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
          iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
          KEGG Genes - Molecular building blocks of life in the genomic space.
          KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
          MEROPS - An information resource for peptidases (also termed proteases, proteinases and proteolytic enzymes) and the proteins that inhibit them.
          modMine - A data warehouse for the modENCODE project
          PDB - An information portal to biological macromolecular structures
          SignaLink - A signaling pathway resource with multi-layered regulatory networks.
          Linkouts
          BioGRID - A database of protein and genetic interactions.
          DroID - A comprehensive database of gene and protein interactions.
          DRSC - Results frm RNAi screens
          FLIGHT - Cell culture data for RNAi and other high-throughput technologies
          FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
          FlyMine - An integrated database for Drosophila genomics
          Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
          InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
          KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
          MIST (genetic) - An integrated Molecular Interaction Database
          MIST (protein-protein) - An integrated Molecular Interaction Database
          Synonyms and Secondary IDs (25)
          Reported As
          Symbol Synonym
          Hh
          (García-Morales et al., 2019, Mirzoyan et al., 2019, Sato et al., 2019, Giordano et al., 2018, Kaur et al., 2018, Lehmann, 2018, Li et al., 2018, Li et al., 2018, Stewart et al., 2018, Walters, 2018, Yu et al., 2018, Amourda and Saunders, 2017, Beer and Wehman, 2017, Chabu et al., 2017, Daniele et al., 2017, Gervais and Bardin, 2017, Hsia et al., 2017, Li et al., 2017, Liu and Jin, 2017, Liu and Jin, 2017, Siddall and Hime, 2017, Suzuki and Sato, 2017, Zhao et al., 2017, Zhou et al., 2017, Beira and Paro, 2016, Czerniak et al., 2016, Dabrowska et al., 2016, Droujinine and Perrimon, 2016, Fried et al., 2016, Jiang et al., 2016, Jiang et al., 2016, Kotov et al., 2016, Lee et al., 2016, Mbodj et al., 2016, Strigini and Leulier, 2016, Yadav et al., 2016, Yasugi and Nishimura, 2016, Bier and De Robertis, 2015, Ciepla et al., 2015, Greenspan et al., 2015, Im et al., 2015, Khaliullina et al., 2015, Lee et al., 2015, Li et al., 2015, Marada et al., 2015, Nagarajan et al., 2015, Oh et al., 2015, Parchure et al., 2015, Simon and Guerrero, 2015, Singh, 2015, Su, 2015, Xie et al., 2015, Xiong et al., 2015, Boekhoff-Falk and Eberl, 2014, Gradilla et al., 2014, Hong and Luo, 2014, Jiang et al., 2014, Kornberg and Roy, 2014, Kuzhandaivel et al., 2014, Lam et al., 2014, Li et al., 2014, Li et al., 2014, Liu et al., 2014, Maier et al., 2014, Pichaud, 2014, Shi et al., 2014, Slattery et al., 2014, Tipping and Perrimon, 2014, Wang et al., 2014, Avanesov and Blair, 2013, Bausek, 2013, Bejsovec, 2013, Briscoe and Thérond, 2013, Chai et al., 2013, Chen and Jiang, 2013, Christiansen et al., 2013, Deshpande et al., 2013, Fan et al., 2013, Gao et al., 2013, Gradilla and Guerrero, 2013, Khuong and Neely, 2013, Lawrence and Casal, 2013, Levayer and Moreno, 2013, Mbodj et al., 2013, Morin-Poulard et al., 2013, Palm et al., 2013, Pastor-Pareja and Xu, 2013, Pepperl et al., 2013, Rana et al., 2013, Sato et al., 2013, Shi et al., 2013, Shim et al., 2013, Verbeni et al., 2013, Yamamoto-Hino and Goto, 2013, Yang et al., 2013, Aikin et al., 2012, Amoyel and Bach, 2012, Ayers et al., 2012, Foronda et al., 2012, Grewal, 2012, Hardy and Resh, 2012, Li et al., 2012, Raftery and Umulis, 2012, Rojas-Ríos et al., 2012, Swarup and Verheyen, 2012, White et al., 2012, Xia et al., 2012, Baker, 2011, Baker and Firth, 2011, Bangi et al., 2011, Brochtrup and Hummel, 2011, Buechling et al., 2011, Chen et al., 2011, Crozatier and Vincent, 2011, Finan et al., 2011, Hadjieconomou et al., 2011, Harris and Ashe, 2011, Kim et al., 2011, Monier et al., 2011, Morata et al., 2011, Roy et al., 2011, Schwank et al., 2011, Shi et al., 2011, Wang et al., 2011, Wang et al., 2011, Wolpert, 2011, Yeung et al., 2011, Zhang et al., 2011, Casali, 2010, Chou et al., 2010, Jia et al., 2010, Li et al., 2010, Raisin et al., 2010, Wang and Hou, 2010, Yan et al., 2010, Zheng et al., 2010, Zhou and Kalderon, 2010, Benítez et al., 2009, Farzan et al., 2009, Gazi et al., 2009, Bornemann et al., 2008, Callejo et al., 2008, Fan and Bergmann, 2008, Franch-Marro et al., 2008, Friggi-Grelin et al., 2008, González et al., 2008, Katanaev et al., 2008, Liu et al., 2008, Su et al., 2008, Ueyama et al., 2008, Vied and Kalderon, 2008, Vyas et al., 2008, Williams et al., 2008, Wojcinski et al., 2008, Zhao and Jiang, 2008, Aikin et al., 2007, Bangi et al., 2007, Bejarano et al., 2007, Callejo et al., 2007, Chien-Hsiang et al., 2007, Escudero and Freeman, 2007, Eugster et al., 2007, Farzan et al., 2007, Firth and Baker, 2007, Gallet et al., 2007, Giuliani et al., 2007, Kugler and Nagel, 2007, Lander, 2007, Makhijani et al., 2007, Molnar et al., 2007, Perrimon and Mathey-Prevot, 2007, Plessis et al., 2007, Reig et al., 2007, Scholler Joulie et al., 2007, Silver et al., 2007, Sun and Deng, 2007, Umemori et al., 2007, Vied and Kalderon, 2007, Wang et al., 2007, Wojcinski et al., 2007, Zhang et al., 2007, Zhao et al., 2007, Beenken and Mohammadi, 2006, Chanana et al., 2006, Croker et al., 2006, De Rivoyre et al., 2006, Fisher and Howie, 2006, Gallet et al., 2006, Goodman et al., 2006, Jia and Jiang, 2006, Joshi et al., 2006, Liu et al., 2006, Maricich and Zoghbi, 2006, McLellan et al., 2006, Ogden et al., 2006, Osterlund and Kogerman, 2006, Sisson et al., 2006, Sisson et al., 2006, Smelkinson and Kalderon, 2006, Wendler et al., 2006, Wilson and Chuang, 2006, Zhang et al., 2006, Zhou et al., 2006, Ziegenhorn et al., 2006, Bovolenta and Marti, 2005, Eldar and Barkai, 2005, Firth and Baker, 2005, Holmgren et al., 2005, Horabin, 2005, Kirkbride, 2005, Ma, 2005, Mehlen et al., 2005, Nybakken et al., 2005, Linder and Deschenes, 2004, Voas and Rebay, 2004, Gonzalez-Gaitan, 2003, Lee and Treisman, 2002, Ma and Beachy, 2002, Gim et al., 2001, Merabet et al., 2001, Selleck et al., 2000, Wang et al., 2000, Zhang and Kalderon, 2000)
          Mir
          anon-WO0134654.19
          anon-WO0182946.19
          hh
          (Bialistoky et al., 2019, Chen, 2019, Copf et al., 2019, Varga et al., 2019, Xu et al., 2019, Aguilar-Hidalgo et al., 2018, Ahaley, 2018, Gáliková and Klepsatel, 2018, Gou et al., 2018, Jia et al., 2018, Jiang et al., 2018, Kittelmann et al., 2018, Lee et al., 2018, Tseng et al., 2018, Aggarwal et al., 2017, Albert and Bökel, 2017, Daniele et al., 2017, Lai et al., 2017, Lu et al., 2017, Muzzopappa et al., 2017, Percival-Smith et al., 2017, Recasens-Alvarez et al., 2017, Takemura and Nakato, 2017, Tian et al., 2017, Transgenic RNAi Project members, 2017-, Wangler et al., 2017, Çiçek et al., 2016, Field et al., 2016, Gene Disruption Project members, 2016-, Iyer et al., 2016, Jin et al., 2016, Li et al., 2016, Moulton and Letsou, 2016, Padash Barmchi et al., 2016, Peng et al., 2016, Willsey et al., 2016, Barr et al., 2015, Liu et al., 2015, Lu et al., 2015, Matsuda et al., 2015, Matsuda et al., 2015, Pasco et al., 2015, Rudolf et al., 2015, Vlachos et al., 2015, Won et al., 2015, Blaquiere et al., 2014, Butí et al., 2014, Camp et al., 2014, Eliazer et al., 2014, Herrera and Morata, 2014, Huang and Kalderon, 2014, Issman-Zecharya and Schuldiner, 2014, Owusu-Ansah and Perrimon, 2014, Sambrani et al., 2014, Aleksic et al., 2013, Baena-Lopez et al., 2013, Bausek, 2013, Chai et al., 2013, Chang et al., 2013, Chauhan et al., 2013, Chen and Jiang, 2013, Da Ros et al., 2013, Deshpande et al., 2013, Ducuing et al., 2013, Ettensohn, 2013, Fossett, 2013, Geisbrecht et al., 2013, Grigorian et al., 2013, Hartman et al., 2013, Huang et al., 2013, Ibrahim et al., 2013, Jin et al., 2013, Kupinski et al., 2013, Li et al., 2013, Nakamura et al., 2013, Palm et al., 2013, Saunders et al., 2013, Singh et al., 2013, Spratford and Kumar, 2013, Tsurui-Nishimura et al., 2013, Webber et al., 2013, Zhang et al., 2013, Aikin et al., 2012, Avanesov et al., 2012, Carroll et al., 2012, Cheng et al., 2012, Cheutin and Cavalli, 2012, Foronda et al., 2012, Hurtado et al., 2012, Jimenez-Sanchez et al., 2012, Krzemien et al., 2012, Mukherjee et al., 2012, Nfonsam et al., 2012, Rojas-Ríos et al., 2012, Sagner et al., 2012, Tokusumi et al., 2012, Bantignies et al., 2011, Callejo et al., 2011, Harterink et al., 2011, Hwang and Rulifson, 2011, Johnson et al., 2011, Johnston et al., 2011, Karim and Moore, 2011, Knox et al., 2011, Marks and Kalderon, 2011, Michaut et al., 2011, Molnar et al., 2011, Ntini and Wimmer, 2011, Ntini and Wimmer, 2011, Pérez et al., 2011, Roy et al., 2011, Schilling et al., 2011, Terriente-Félix et al., 2011, Toku et al., 2011, Watson et al., 2011, Yuva-Aydemir et al., 2011, Zhang et al., 2011, Ayers et al., 2010, Baig et al., 2010, Bergantiños et al., 2010, Biehs et al., 2010, Chang et al., 2010, Cheng et al., 2010, Dilks and DiNardo, 2010, Hartman et al., 2010, Irons et al., 2010, Klein et al., 2010, Liu et al., 2010, Lopes and Casares, 2010, Maurel-Zaffran et al., 2010, Pospisilik et al., 2010, Salzer and Kumar, 2010, Sato et al., 2010, Schwartz et al., 2010, Seong et al., 2010, Smulders-Srinivasan et al., 2010, Subramanian and Gadgil, 2010, Terriente-Félix et al., 2010, Tokhunts et al., 2010, Wang et al., 2010, Williams et al., 2010, Yavari et al., 2010, Zheng et al., 2010, Zhou and Kalderon, 2010, Baker et al., 2009, Bejarano and Milán, 2009, Blanco et al., 2009, Chaves et al., 2009, Christensen et al., 2009.5.6, Deshpande et al., 2009, Eivers et al., 2009, Foronda et al., 2009, Gazi et al., 2009, González et al., 2009, Gutierrez-Aviño et al., 2009, Jia et al., 2009, Julius et al., 2009, Khaliullina et al., 2009, Landsberg et al., 2009, Langmead and Jha, 2009, May and Schiek, 2009, Mulinari and Häcker, 2009, Nahmad and Stathopoulos, 2009, Renault et al., 2009, Schuettengruber et al., 2009, Southall and Brand, 2009, Venken et al., 2009, Vied and Kalderon, 2009, Wang and Huang, 2009, Bornemann et al., 2008, Brás-Pereira and Casares, 2008, Casso et al., 2008, Casso et al., 2008, Chaves and Albert, 2008, Chen et al., 2008, Christensen et al., 2008.9.29, Christensen et al., 2008.9.29, Fan and Bergmann, 2008, Farzan et al., 2008, Gallet et al., 2008, Hallson et al., 2008, Larsen et al., 2008, Lim et al., 2008, McLellan et al., 2008, Melicharek et al., 2008, Ogden et al., 2008, Sánchez et al., 2008, Sato et al., 2008, Schlichting and Dahmann, 2008, Takashima et al., 2008, Vincent et al., 2008, Vyas et al., 2008, Wang and Price, 2008, Wang et al., 2008, Zhao et al., 2008, Bejarano et al., 2007, Bejarano et al., 2007, Beltran et al., 2007, Bras-Pereira and Casares, 2007, Casso et al., 2007, Chanana et al., 2007, Chien-Hsiang et al., 2007, DasGupta et al., 2007, Deshpande et al., 2007, de Velasco et al., 2007, Lechner et al., 2007, Lindner et al., 2007, Lindner et al., 2007, Liu et al., 2007, Maeda et al., 2007, Magalhaes et al., 2007, Mandal et al., 2007, Molnar et al., 2007, Ntini E and Wimmer, 2007, Ou et al., 2007, Pfleger et al., 2007, Pichaud et al., 2007, Sakurai et al., 2007, Sandmann et al., 2007, Smelkinson et al., 2007, Song et al., 2007, Sprecher et al., 2007, Su et al., 2007, Tountas and Fortini, 2007, Walthall et al., 2007, Bras-Pereira et al., 2006, Callejo et al., 2006, Chanut-Delalande et al., 2006, Chu et al., 2006, Colosimo and Tolwinski, 2006, D'Costa et al., 2006, de Velasco et al., 2006, Fraser, 2006, Friedrich, 2006, Guichard et al., 2006, Jones et al., 2006, Kent et al., 2006, Lu et al., 2006, Mahoney et al., 2006, Martin-Lanneree et al., 2006, Molnar et al., 2006, Nystul and Spradling, 2006, Price et al., 2006, Ramos and Mohler, 2006, Smelkinson and Kalderon, 2006, Suh et al., 2006, Umetsu et al., 2006, Vrailas and Moses, 2006, Wheeler et al., 2006, Yao et al., 2006, Yasunaga et al., 2006, Akimoto et al., 2005, Besse et al., 2005, Briscoe and Therond, 2005, Chanas and Maschat, 2005, Chotard et al., 2005, Dawber et al., 2005, Deshpande and Schedl, 2005, Deshpande and Schedl, 2005, Glazov et al., 2005, Glise et al., 2005, Gorfinkiel et al., 2005, Ishii, 2005, Peel et al., 2005, Roederer et al., 2005, Rogers et al., 2005, Torroja et al., 2005, Xie et al., 2005, Zhang et al., 2005, Cheesman et al., 2004, Wang and Struhl, 2004, Monnier et al., 2002, Hayashi and Murakami, 2001, Birdsall et al., 2000, Emerald and Shashidhara, 2000, Levine et al., 1997, Freeland and Kuhn, 1996)
          l(3)neo57
          Name Synonyms
          Hedgehog
          (Xu et al., 2019, Roelink, 2018, Schwartz and Rhiner, 2018, Yu et al., 2018, Beer and Wehman, 2017, Lefebvre et al., 2017, Takemura and Nakato, 2017, Zhao et al., 2017, Czerniak et al., 2016, Fried et al., 2016, Parsons and Foley, 2016, Buchon and Osman, 2015, Irvine and Harvey, 2015, Lu et al., 2015, Matsuda et al., 2015, Parchure et al., 2015, Rudolf et al., 2015, Vlisidou and Wood, 2015, Jones and Srivastava, 2014, Shi et al., 2014, Avanesov and Blair, 2013, Chai et al., 2013, Christiansen et al., 2013, Fan et al., 2013, Gradilla and Guerrero, 2013, Hartman et al., 2013, Lawrence and Casal, 2013, Levayer and Moreno, 2013, Morin-Poulard et al., 2013, Palm et al., 2013, Pepperl et al., 2013, Shi et al., 2013, Solis et al., 2013, Verbeni et al., 2013, Weasner and Kumar, 2013, Yamamoto-Hino and Goto, 2013, Yang et al., 2013, Amoyel and Bach, 2012, Cheng et al., 2012, Fan et al., 2012, Krzemien et al., 2012, Li et al., 2012, Rincon-Limas et al., 2012, Rojas-Ríos et al., 2012, Sagner et al., 2012, White et al., 2012, Dahmann et al., 2011, Harris and Ashe, 2011, Harterink et al., 2011, Ingham et al., 2011, Juarez et al., 2011, Molnar et al., 2011, Monier et al., 2011, Niwa and Niwa, 2011, Pocha et al., 2011, Roy et al., 2011, Schilling et al., 2011, Shi et al., 2011, Wang et al., 2011, Casali, 2010, Chang et al., 2010, Chou et al., 2010, Hartman et al., 2010, Irons et al., 2010, Jia et al., 2010, Maurel-Zaffran et al., 2010, Raisin et al., 2010, Terriente-Félix et al., 2010, Yan et al., 2010, Zheng et al., 2010, Zhou and Kalderon, 2010, Baker et al., 2009, Buchon et al., 2009, Farzan et al., 2009, Gazi et al., 2009, Khaliullina et al., 2009, Renault et al., 2009, Baudot et al., 2008, Bornemann et al., 2008, Casso et al., 2008, Fan and Bergmann, 2008, Farzan et al., 2008, Franch-Marro et al., 2008, Gallet et al., 2008, Legent et al., 2008, Lim et al., 2008, Liu et al., 2008, Lorigan et al., 2008, Ming et al., 2008, Vied and Kalderon, 2008, Williams et al., 2008, Zhao and Jiang, 2008, Bejarano et al., 2007, Bulanin and Orenic, 2007, Chien-Hsiang et al., 2007, Claret et al., 2007, Coudreuse and Korswagen, 2007, Dahmann and Schlichting, 2007, Escudero and Freeman, 2007, Escudero et al., 2007, Eugster et al., 2007, Giuliani et al., 2007, Kerszberg and Wolpert, 2007, Lindner et al., 2007, Liu et al., 2007, Mandal et al., 2007, Molnar et al., 2007, Payre et al., 2007, Plessis et al., 2007, Plessis et al., 2007, Reig et al., 2007, Sanial and Plessis, 2007, Schlichting and Dahmann, 2007, Schwartz and Pirrotta, 2007, Su et al., 2007, Vied et al., 2007, Walthall et al., 2007, Wojcinski et al., 2007, Zhao et al., 2007, Zinzen and Papatsenko, 2007, Arbouzova and Zeidler, 2006, Beach et al., 2006, Bossing and Brand, 2006, Chu et al., 2006, Colosimo and Tolwinski, 2006, Friedrich, 2006, Gallet et al., 2006, Goodman et al., 2006, Maurange et al., 2006, McLellan et al., 2006, Niki et al., 2006, Ogden et al., 2006, Panakova and Eaton, 2006, Sisson et al., 2006, Tolhuis et al., 2006, Vrailas et al., 2006, Dawber et al., 2005, Horabin, 2005, Pallavi and Shashidhara, 2005, Strigini, 2005, Torroja et al., 2005, Lawrence, 2004, Linder and Deschenes, 2004, Voas and Rebay, 2004, Monnier et al., 2002, Gim et al., 2001)
          Mirabile
          Moonrat
          bar-on-3
          hedgehog
          (Courcoubetis et al., 2019, García-Morales et al., 2019, Meltzer et al., 2019, Li et al., 2017, Wangler et al., 2017, Çiçek et al., 2016, Wieschaus and Nüsslein-Volhard, 2016, Ghimire and Kim, 2015, Hallier et al., 2015, Matsuda et al., 2015, Moncrieff et al., 2015, Oh et al., 2015, Pasco et al., 2015, Simon and Guerrero, 2015, Xiong et al., 2015, Gradilla et al., 2014, Kim et al., 2014, Lam et al., 2014, Owusu-Ansah and Perrimon, 2014, Wang et al., 2014, Chauhan et al., 2013, Da Ros et al., 2013, Deshpande et al., 2013, Geisbrecht et al., 2013, Ibrahim et al., 2013, Khuong and Neely, 2013, Marianes and Spradling, 2013, Marques-Pita and Rocha, 2013, Rana et al., 2013, Spratford and Kumar, 2013, Zhan et al., 2013, Matunis et al., 2012, Tokusumi et al., 2012, Zoller and Schulz, 2012, Eliazer and Buszczak, 2011, Finan et al., 2011, Johnson et al., 2011, Johnston et al., 2011, Ntini and Wimmer, 2011, Ntini and Wimmer, 2011, Pérez et al., 2011, Terriente-Félix et al., 2011, Baig et al., 2010, Biehs et al., 2010, Cheng et al., 2010, Dilks and DiNardo, 2010, Liu et al., 2010, Lopes and Casares, 2010, Salzer and Kumar, 2010, Schwartz et al., 2010, Tokusumi et al., 2010, Williams et al., 2010, Yavari et al., 2010, Eivers et al., 2009, Gutierrez-Aviño et al., 2009, Jia et al., 2009, Martinez et al., 2009, Mulinari and Häcker, 2009, Dansereau and Lasko, 2008, Ishihara and Shibata, 2008, McLellan et al., 2008, Melicharek et al., 2008, Mulinari et al., 2008, Schlichting and Dahmann, 2008, Takashima et al., 2008, Bejarano et al., 2007, Bras-Pereira and Casares, 2007, Chanana et al., 2007, DasGupta et al., 2007, Lander, 2007, Lechner et al., 2007, Maeda et al., 2007, Mulinari et al., 2007, Ou et al., 2007, Song et al., 2007, Thompson et al., 2007, Tountas and Fortini, 2007, de Velasco et al., 2006, Fraser, 2006, Hashimoto and Yamaguchi, 2006, Muller and Kassis, 2006, Nystul and Spradling, 2006, Price et al., 2006, Vrailas and Moses, 2006, Wheeler et al., 2006, Akimoto et al., 2005, Besse et al., 2005, Glazov et al., 2005, Kruger, 2005, Roederer et al., 2005, Rogers et al., 2005, Thomas, 2005, Hime et al., 2004, Gonzalez-Reyes, 2003, Birdsall et al., 2000, Gellon et al., 1997)
          Secondary FlyBase IDs
          • FBgn0000159
          • FBgn0001191
          • FBgn0002748
          • FBgn0002793
          • FBgn0011486
          • FBgn0011487
          • FBgn0044801
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          References (1,902)