FB2026_02 , released June 18, 2026
Gene: Dmel\smo
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General Information
Symbol
Dmel\smo
Species
D. melanogaster
Name
smoothened
Annotation Symbol
CG11561
Feature Type
protein_coding_gene
FlyBase ID
FBgn0003444
Gene Model Status
Current
Stock Availability
41 publicly available
Gene Summary
smoothened (smo) encodes a critical component of the Hedgehog signaling pathway. It is regulated by phosphorylation, dimerization, and cell-surface accumulation upon Hedgehog stimulation. The product of smo is also regulated by ubiquitin and other small molecules such as phospholipids. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
All Summaries
Also Known As

dSmo, smooth

Key Links
Genomic Location
Cytogenetic map
21B7-21B7
Sequence location
Recombination map
2-0.5
RefSeq locus
NT_033779 REGION:277582..282167
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (37 terms)
Molecular Function (8 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
enables G protein-coupled receptor activity
inferred from direct assay
(Ogden et al., 2008)
enables phosphatidic acid binding
inferred from direct assay
(Zhang et al., 2023)
enables phosphatidylinositol-4-phosphate binding
inferred from direct assay
(Jiang et al., 2016)
enables protein binding
inferred from physical interaction with FLYBASE:cos; FB:FBgn0000352
(Fan et al., 2012, Jia et al., 2003, Ogden et al., 2003)
inferred from physical interaction with UniProtKB:O16844
(Zhang et al., 2005)
inferred from physical interaction with FLYBASE:Sxl; FB:FBgn0264270
(Walthall et al., 2007)
enables protein homodimerization activity
inferred from direct assay
(Shi et al., 2011)
enables protein kinase binding
inferred from physical interaction with FLYBASE:fu; FB:FBgn0001079
(Zhang et al., 2011)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables patched binding
inferred from biological aspect of ancestor with PANTHER:PTN000885245
(GO Reference Genome Project, 2011-)
enables transmembrane signaling receptor activity
inferred from electronic annotation with InterPro:IPR017981
(InterPro Project Members, 2004-)
Biological Process (23 terms)
Terms Based on Experimental Evidence (17 terms)
CV Term
Evidence
References
involved_in adenylate cyclase-inhibiting G protein-coupled receptor signaling pathway
inferred from mutant phenotype
(Ogden et al., 2008)
involved_in blastoderm segmentation
inferred from mutant phenotype
(van den Heuvel and Ingham, 1996)
involved_in Bolwig's organ morphogenesis
inferred from mutant phenotype
(Suzuki and Saigo, 2000)
involved_in compound eye morphogenesis
inferred from mutant phenotype
(Kent et al., 2006)
involved_in eye-antennal disc morphogenesis
inferred from genetic interaction with FLYBASE:ptc; FB:FBgn0003892
(Shyamala and Bhat, 2002)
involved_in follicle cell of egg chamber development
inferred from genetic interaction with FLYBASE:dsh; FB:FBgn0000499
(Dai et al., 2017)
inferred from mutant phenotype
(Kirilly et al., 2005)
involved_in imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Chen et al., 2010)
involved_in mucosal immune response
inferred from mutant phenotype
(Lee et al., 2015)
involved_in negative regulation of G1/S transition of mitotic cell cycle
inferred from mutant phenotype
(Firth and Baker, 2005)
acts_upstream_of positive regulation of gene expression
inferred from mutant phenotype
(Yang et al., 2022)
involved_in positive regulation of protein localization to plasma membrane
inferred from direct assay
(Claret et al., 2007)
involved_in regulation of apoptotic process
inferred from genetic interaction with FLYBASE:hid; FB:FBgn0003997
(Christiansen et al., 2013)
involved_in regulation of mitotic cell cycle
inferred from mutant phenotype
(Zhang and Kalderon, 2001)
involved_in regulation of stem cell differentiation
inferred from mutant phenotype
(Huang et al., 2017)
involved_in segment polarity determination
inferred from mutant phenotype
(van den Heuvel and Ingham, 1996)
involved_in smoothened signaling pathway
inferred from mutant phenotype
(He et al., 2022, Yang et al., 2022, Han et al., 2019, Giordano et al., 2018, Sanial et al., 2017, Shi et al., 2014, Cheng et al., 2012, Zhou and Kalderon, 2011, Chen et al., 2010, Cheng et al., 2010, Raisin et al., 2010, Liu et al., 2007, Ogden et al., 2006, Apionishev et al., 2005, Jia et al., 2004, Zhang et al., 2004, Jia et al., 2003, Ogden et al., 2003, Ruel et al., 2003, Chen and Struhl, 1996)
inferred from genetic interaction with FLYBASE:ci; FB:FBgn0004859
(Oh et al., 2015, Smelkinson and Kalderon, 2006)
inferred from direct assay
(Gonçalves Antunes et al., 2022, Li et al., 2014, Ranieri et al., 2014, Carroll et al., 2012, Fan et al., 2012, Ranieri et al., 2012, Shi et al., 2011, Zhang et al., 2011, Zhao et al., 2007, Zhang et al., 2005, Jia et al., 2004, Lum et al., 2003)
inferred from direct assay
(Ogden et al., 2008, Denef et al., 2000)
inferred from genetic interaction with FLYBASE:fu; FB:FBgn0001079
(Malpel et al., 2007)
involved_in somatic stem cell population maintenance
inferred from mutant phenotype
(Kirilly et al., 2005)
Terms Based on Predictions or Assertions (7 terms)
CV Term
Evidence
References
involved_in anterior/posterior lineage restriction, imaginal disc
traceable author statement
(Milan and Cohen, 2000)
involved_in cell surface receptor signaling pathway
inferred from electronic annotation with InterPro:IPR000539, InterPro:IPR017981
(InterPro Project Members, 2004-)
involved_in central nervous system development
inferred from biological aspect of ancestor with PANTHER:PTN000885245
(GO Reference Genome Project, 2011-)
involved_in commissural neuron axon guidance
inferred from biological aspect of ancestor with PANTHER:PTN000885245
(GO Reference Genome Project, 2011-)
involved_in pattern specification process
inferred from biological aspect of ancestor with PANTHER:PTN000885245
(GO Reference Genome Project, 2011-)
involved_in smoothened signaling pathway
inferred from biological aspect of ancestor with PANTHER:PTN000885245
(GO Reference Genome Project, 2011-)
involved_in wing disc anterior/posterior pattern formation
traceable author statement
(McNeill, 2000)
Cellular Component (6 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
colocalizes_with axon
inferred from direct assay
(Kuzhandaivel et al., 2014)
located_in cilium
inferred from direct assay
(Kuzhandaivel et al., 2014)
located_in dendrite
inferred from direct assay
(Kuzhandaivel et al., 2014)
located_in neuronal cell body
inferred from direct assay
(Kuzhandaivel et al., 2014)
located_in plasma membrane
inferred from mutant phenotype
(Cheng et al., 2010)
inferred from direct assay
(Carroll et al., 2012, Fan et al., 2012, Zhao et al., 2007, Denef et al., 2000)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
is_active_in cilium
inferred from biological aspect of ancestor with PANTHER:PTN000885245
(GO Reference Genome Project, 2011-)
is_active_in dendrite
inferred from biological aspect of ancestor with PANTHER:PTN000885245
(GO Reference Genome Project, 2011-)
located_in membrane
inferred from electronic annotation with InterPro:IPR000539, InterPro:IPR017981
(InterPro Project Members, 2004-)
is_active_in plasma membrane
inferred from biological aspect of ancestor with PANTHER:PTN000139394
(GO Reference Genome Project, 2011-)
located_in plasma membrane
inferred from experiment
(Jia et al., 2004)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
Belongs to the G-protein coupled receptor Fz/Smo family. (P91682)
Protein Signatures (InterPro)
Summaries
Gene Snapshot
smoothened (smo) encodes a critical component of the Hedgehog signaling pathway. It is regulated by phosphorylation, dimerization, and cell-surface accumulation upon Hedgehog stimulation. The product of smo is also regulated by ubiquitin and other small molecules such as phospholipids. [Date last reviewed: 2019-03-14]
Gene Group (FlyBase)
SMOOTHENED-TYPE RECEPTORS -
smo is a Class F GPCR related to Frizzled GPCRs. It is a component of the hedgehog (hh) signaling pathway. smo is believed to signal via G protein independent and Gi-coupled pathways. (Adapted from FBrf0221081 and FBrf0201812).
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).
Protein Function (UniProtKB)
Segment polarity protein required for correct patterning of every segment. G protein-coupled receptor which associates with the patched protein (ptc) to transduce the hedgehog (hh) signal through the activation of an inhibitory G-protein. In the absence of hh, ptc represses the constitutive signaling activity of smo through fused (fu). Essential component of a hh-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).
(UniProt, P91682)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
smo: smoothened
Embryonic lethal. All denticles in abdominal segments point posteriorly. At 18 naked cuticle deleted and denticle belts of adjacent segments fused and locally arranged as mirror-image duplications.
Summary (Interactive Fly)

seven-pass transmembrane protein - the reception and transduction of the HH signal is mediated by its receptor Patched and by Smoothened - PTC and HH control SMO by regulating its stability, trafficking, and phosphorylation - SMO in turn interacts directly with Fused and Costal2, which interact with each other and with Cubitus interruptus in an intracellular Hedgehog transducing complex

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

Please see the JBrowse view of Dmel\smo 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 P91682)

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
PDB - An information portal to biological macromolecular structures
Comments on Gene Model

Gene model reviewed during 5.44

Gene model reviewed during 5.52

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
smo-RA
FBtr0078129
NM_078719
4008
1036
Additional Transcript Data and Comments
Reported size (kB)

4.2 (northern blot)

(Alcedo et al., 1996)

4.1 (northern blot)

(van den Heuvel and Ingham, 1996)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
smo-PA
FBpp0077788
116.6
1036
8.95
P91682
NP_523443
AAF51518
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)

1036 (aa)

(Alcedo et al., 1996)
Comments
External Data
Subunit Structure (UniProtKB)

Interacts with cos.

(UniProt, P91682)
Post Translational Modification

Phosphorylation by CkIalpha and PKA regulates smo accumulation at the cell surface and its signaling activity in response to hh.

(UniProt, P91682)
Crossreferences
InterPro - A database of protein families, domains and functional sites
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\smo 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).

-0.66

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
epithelial cell
• cytoplasm | apical

Comment: clustered apical distribution

(Lecuyer et al., 2007)
epithelium
• cytoplasm | apical
(Lecuyer et al., 2007)
embryonic stage 1 -- 3
pole plasm

Comment: maternally deposited

(Fisher et al., 2012)
organism
• pole plasm

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
central brain anlage in statu nascendi
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
antennal anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

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

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)
visual anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)
embryonic stage 7 -- 8
antennal anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
central brain anlage

Comment: reported as procephalic ectoderm anlage

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

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
visual anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
dorsal ectoderm
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
embryonic stage 9 -- 10
antennal primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
central brain primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
visual primordium

Comment: reported as procephalic ectoderm primordium

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

Comment: reported as procephalic ectoderm primordium

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

Comment: reported as procephalic ectoderm primordium

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

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
inclusive hindgut primordium
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
embryonic stage 11
visual primordium | restricted
(Biehs et al., 2010)
embryonic stage 11 -- 12
antennal primordium

Comment: reported as procephalon primordium

(Fisher et al., 2012)
central brain primordium

Comment: reported as procephalon primordium

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

Comment: reported as procephalon primordium

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

Comment: reported as procephalon primordium

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

Comment: reported as procephalon primordium

(Fisher et al., 2012)
visual primordium

Comment: reported as procephalon primordium

(Fisher et al., 2012)
pars intercerebralis primordium
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic dorsal epidermis
(Fisher et al., 2012)
wandering third instar larval stage
wing disc
(Molnar et al., 2007)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage & larval stage & adult stage
(van den Heuvel and Ingham, 1996)
RT-PCR
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
embryonic/larval fat body
(Suh et al., 2006)
Additional Descriptive Data

smo is present in both the posterior optic lobe and adjacent anterior optic lobe cells in mid stage 11 embryos.

(Biehs et al., 2010)

smo transcript is expressed throughout the wing disc.

(Molnar et al., 2007)

smo transcripts are expressed at all developmental stages.

(Alcedo et al., 1996)

smo transcripts are detected in embryos, larvae, and pupae on northern blots. They are most abundant in early embryos and adult females.

(van den Heuvel and Ingham, 1996)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
wing disc
(Gao et al., 2013, Cheng et al., 2010)
wing disc posterior compartment | intense
(Gao et al., 2013)
wandering third instar larval stage
wing disc anterior compartment
(Molnar et al., 2007)
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
day 5 of adulthood | male -- day 7 of adulthood | male
spermatozoon
(Wasbrough et al., 2010)
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
wing disc
(Cheng et al., 2010)
Additional Descriptive Data

smo protein is localized to the anterior compartment of the wing disc; however, smo transcipt is distributed through the entire wing disc.

(Molnar et al., 2007)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
colocalizes_with axon
inferred from direct assay
(Kuzhandaivel et al., 2014)
located_in cilium
inferred from direct assay
(Kuzhandaivel et al., 2014)
located_in dendrite
inferred from direct assay
(Kuzhandaivel et al., 2014)
located_in neuronal cell body
inferred from direct assay
(Kuzhandaivel et al., 2014)
located_in plasma membrane
inferred from mutant phenotype
(Cheng et al., 2010)
inferred from direct assay
(Carroll et al., 2012, Fan et al., 2012, Zhao et al., 2007, Denef et al., 2000)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\smo in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 9-10
  • Stages(s) 11-12
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 24 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 252 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of smo
Transgenic constructs containing regulatory region of smo
Aberrations (Deficiencies and Duplications) ( 7 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal, with Scer\GAL4btl.PS
lethal (with Df(2L)BSC454)
lethal (with smo1K)
lethal (with smo2A)
lethal (with smo3)
lethal (with smo3J)
lethal (with smo4C2)
lethal (with smo4DI)
lethal (with smo4F4)
lethal (with smoIA3)
lethal | recessive
lethal | recessive | cold sensitive
lethal - all die before end of P-stage, with Scer\GAL471B
lethal - all die before end of P-stage, with Scer\GAL4Bx-MS1096
lethal - all die before end of pupal stage, with Scer\GAL4ptc-559.1
lethal - all die during embryonic stage, with Scer\GAL4Act5C.PI
lethal - all die during embryonic stage | recessive
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4C-765
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4tin.CΔ4
Sterility
Allele
fertile, with Scer\GAL4C-765
Other Phenotypes
Allele
abnormal immune response, with Scer\GAL4NP0001
abnormal mitotic cell cycle | cell autonomous | somatic clone
abnormal mitotic cell cycle | cell non-autonomous | somatic clone
abnormal mitotic cell cycle | somatic clone
abnormal size, with Scer\GAL4MD-638
abnormal size | adult stage, with Scer\GAL4nub.PU
abnormal size | somatic clone
decreased cell number, with Scer\GAL4C587
decreased cell number | larval stage, with Scer\GAL4btl.PS
decreased cell number | oogenesis, with Scer\GAL4C587
decreased cell number | oogenesis | somatic clone
decreased cell number | somatic clone
decreased cell number | somatic clone, with Scer\GAL4αTub84B.PP
decreased occurrence of cell division | larval stage, with Scer\GAL4btl.PS
decreased occurrence of cell division | larval stage | somatic clone
decreased occurrence of cell division | somatic clone | third instar larval stage
decreased size | adult stage, with Scer\GAL4btl.PS
decreased size | larval stage, with Scer\GAL4btl.PS
increased cell death | somatic clone
increased cell number, with Scer\GAL4C587
increased cell number, with Scer\GAL4GMR25A11
increased cell number | oogenesis, with Scer\GAL4C587
increased occurrence of cell division | oogenesis | somatic clone
neoplasia | adult stage | female, with Scer\GAL4C587
segment polarity phenotype | cold sensitive
segment polarity phenotype | recessive | cold sensitive
short lived, with Scer\GAL4NP0001
some die during embryonic stage, with Scer\GAL4Act5C.PI
some die during embryonic stage | recessive
some die during pharate adult stage, with Scer\GAL471B
some die during pharate adult stage, with Scer\GAL4Bx-MS1096
some die during pupal stage, with Scer\GAL4ptc-559.1
visible
visible, with Scer\GAL471B
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4C-765
visible, with Scer\GAL4C-765, smo3
visible, with Scer\GAL4C-765, smo5A.UAS.Tag:FLAG
visible, with Scer\GAL4MD-638
visible, with Scer\GAL4Mef2.PR
visible, with Scer\GAL4ptc-559.1
visible, with Scer\GAL4ptc-559.1, smo3
visible, with Scer\GAL4ptc-559.1, smoRNAi.UAS.WIZ
visible, with Scer\GAL4ptc.PU
visible, with Scer\GAL4T80
visible | adult stage, with Scer\GAL4C-765
visible | adult stage, with Scer\GAL4nub.PU
visible | recessive | somatic clone
visible | somatic clone
wild-type
wild-type, with Scer\GAL4h-1J3
wild-type, with Scer\GAL4prd.RG1
Phenotype manifest in
Allele
abdominal 1 ventral denticle belt
abdominal 2 ventral denticle belt
abdominal 3 ventral denticle belt
abdominal 4 ventral denticle belt
abdominal 5 ventral denticle belt
abdominal 6 ventral denticle belt
abdominal 7 ventral denticle belt
abdominal 8 ventral denticle belt
abdominal segment 10 anterior compartment
abdominal segment 11 anterior compartment
abdominal tergite 8 | somatic clone
abdominal tergite | anterior | somatic clone
adult abdominal segment anterior compartment
adult cuticle | somatic clone
adult head | somatic clone
adult head capsule | somatic clone
adult thorax | somatic clone
air sac primordium | larval stage, with Scer\GAL4btl.PS
anterior-posterior compartment boundary of the wing blade, with Scer\GAL4ptc.PU
anterior-posterior compartment boundary of the wing disc
Bolwig organ | germline clone
campaniform sensillum of anterior crossvein, with Scer\GAL4Bx-MS1096
cercus | somatic clone
costal wing cell, with Scer\GAL4Bx-MS1096
crossvein, with Scer\GAL4Bx-MS1096
crossvein | ectopic, with Scer\GAL471B
crossvein | ectopic, with Scer\GAL4Bx-MS1096
crossvein | ectopic, with Scer\GAL4C-765
cuticle
cytoneme | larval stage, with Scer\GAL4btl.PS
denticle | germline clone | increased number
dorsal air sac | adult stage, with Scer\GAL4btl.PS
double row of wing sensilla | ectopic | somatic clone
egg chamber, with Scer\GAL4c306
egg chamber | somatic clone
embryonic/first instar larval cuticle
embryonic/first instar larval cuticle | cold sensitive
embryonic/first instar larval cuticle | germline clone
embryonic/larval dorsal tracheal branch, with Scer\GAL4btl.PS
embryonic epidermis
embryonic epidermis | cold sensitive
embryonic epidermis | germline clone
epandrium | somatic clone
epiproct | somatic clone
escort cell | decreased number, with Scer\GAL4C587
eye, with Scer\GAL4lz-gal4
eye | somatic clone
eye disc | cell autonomous | somatic clone
eye disc | cell non-autonomous | somatic clone
eye disc | somatic clone
eye disc | somatic clone | third instar larval stage
eye disc morphogenetic furrow
eye disc morphogenetic furrow | somatic clone
female germline cell | increased number, with Scer\GAL4C587
first posterior wing cell, with Scer\GAL4Bx-MS1096
first posterior wing cell, with Scer\GAL4C-765
first posterior wing cell, with Scer\GAL4C-765, smo3
first posterior wing cell, with Scer\GAL4C-765, smo5A.UAS.Tag:FLAG
follicle cell | somatic clone
follicle stem cell | somatic clone
germarium, with Scer\GAL4C587
germarium, with Scer\GAL4GMR25A11
germarium region 3 | somatic clone
germline cell | embryonic stage | germline clone
hypandrial bristle | somatic clone
hypogynium | somatic clone
imaginal disc anterior compartment, with Scer\GAL4MD-638
lamina & neuron | precursor | somatic clone
larval tracheal system, with Scer\GAL4btl.PS
leg | somatic clone
maxillary palp | somatic clone
maxillary palp olfactory receptor neuron | cell non-autonomous | somatic clone
medial ocellus | somatic clone | increased number
microchaeta
microchaeta | ectopic | somatic clone
morphogenetic furrow & nucleus | somatic clone
ocellus | somatic clone
ommatidial precursor cluster
ommatidial precursor cluster, with Scer\GAL4lz-gal4
ovariole, with Scer\GAL4c306
ovariole | somatic clone
photoreceptor cell & neuron
photoreceptor cell R8 & eye disc | somatic clone
photoreceptor cell R8 | precursor
photoreceptor neuron
photoreceptor neuron | cell non-autonomous | somatic clone
photoreceptor neuron | somatic clone
primordial germ cell | embryonic stage | maternal effect, with Scer\GAL4Tub.PU
scutellum | somatic clone
spectrosome, with Scer\GAL4C587
stalk follicle cell | somatic clone
submarginal wing cell, with Scer\GAL4Bx-MS1096
surstylus | somatic clone
tergite anterior compartment
third segment of antenna | somatic clone
ventral denticle belt
wing
wing, with Scer\GAL471B
wing, with Scer\GAL4Bx-MS1096
wing, with Scer\GAL4C-765
wing, with Scer\GAL4MD-638
wing, with Scer\GAL4Mef2.PR
wing, with Scer\GAL4nub.PU
wing | somatic clone
wing blade, with Scer\GAL4C-765
wing blade, with Scer\GAL4MD-638
wing blade, with Scer\GAL4ptc.PU
wing blade | anterior, with Scer\GAL4C-765
wing blade anterior compartment
wing blade anterior compartment, with Scer\GAL4en-e16E
wing blade posterior compartment | somatic clone
wing cell, with Scer\GAL471B
wing cell, with Scer\GAL4Bx-MS1096
wing disc | larval stage, with Scer\GAL41151
wing disc | somatic clone
wing disc | third instar larval stage, with Scer\GAL4MD-638
wing margin | somatic clone
wing pouch | somatic clone
wing vein
wing vein, with Scer\GAL4C-765
wing vein | ectopic
wing vein | ectopic, with Scer\GAL471B
wing vein | ectopic, with Scer\GAL4Bx-MS1096
wing vein | ectopic, with Scer\GAL4C-765
wing vein | ectopic | somatic clone
wing vein L1, with Scer\GAL4Bx-MS1096
wing vein L2, with Scer\GAL4MD-638
wing vein L3, with Scer\GAL471B
wing vein L3, with Scer\GAL4Bx-MS1096
wing vein L3, with Scer\GAL4C-765
wing vein L3, with Scer\GAL4C-765, smo3
wing vein L3, with Scer\GAL4C-765, smo5A.UAS.Tag:FLAG
wing vein L3, with Scer\GAL4MD-638
wing vein L3, with Scer\GAL4ptc-559.1
wing vein L3, with Scer\GAL4ptc-559.1, smo3
wing vein L3, with Scer\GAL4ptc-559.1, smoRNAi.UAS.WIZ
wing vein L3, with Scer\GAL4T80
wing vein L3 | proximal, with Scer\GAL4ptc-559.1
wing vein L4, with Scer\GAL4Bx-MS1096
wing vein L4, with Scer\GAL4C-765
wing vein L4, with Scer\GAL4C-765, smo3
wing vein L4, with Scer\GAL4C-765, smo5A.UAS.Tag:FLAG
wing vein L4, with Scer\GAL4ptc-559.1
wing vein L4, with Scer\GAL4ptc-559.1, smo3
wing vein L4, with Scer\GAL4ptc-559.1, smoRNAi.UAS.WIZ
wing vein L4, with Scer\GAL4T80
wing vein L4 | ectopic | somatic clone
wing vein L4 | proximal, with Scer\GAL4ptc-559.1
wing vein L4 | somatic clone
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (24)
Hsap\SMO
13 of 14
Yes
Yes
Hsap\FZD4
3 of 14
No
No
Hsap\FZD5
3 of 14
No
No
Hsap\FZD8
3 of 14
No
No
Hsap\FZD9
3 of 14
No
No
Hsap\FZD10
3 of 14
No
No
Hsap\CORIN
2 of 14
No
No
Hsap\FRZB
2 of 14
No
No
Hsap\FZD1
2 of 14
No
No
Hsap\FZD2
2 of 14
No
No
Hsap\FZD3
2 of 14
No
No
Hsap\FZD6
2 of 14
No
No
Hsap\FZD7
2 of 14
No
No
3  
Hsap\ADGRB1
1 of 14
No
No
Hsap\ADGRB2
1 of 14
No
No
Hsap\ADGRG4
1 of 14
No
No
Hsap\ADGRL3
1 of 14
No
No
Hsap\COL18A1
1 of 14
No
No
Hsap\CPZ
1 of 14
No
No
Hsap\MFRP
1 of 14
No
No
Hsap\SFRP1
1 of 14
No
No
Hsap\SFRP2
1 of 14
No
No
Hsap\SFRP4
1 of 14
No
No
Hsap\SFRP5
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (20)
Rnor\Smo
13 of 14
Yes
Yes
Rnor\Fzd4
3 of 14
No
No
Rnor\Fzd8
3 of 14
No
No
Rnor\Fzd9
3 of 14
No
No
Rnor\Corin
2 of 14
No
No
Rnor\Frzb
2 of 14
No
No
Rnor\Fzd1
2 of 14
No
No
Rnor\Fzd2
2 of 14
No
No
Rnor\Fzd3
2 of 14
No
No
Rnor\Fzd5
2 of 14
No
No
Rnor\Fzd6
2 of 14
No
No
Rnor\Fzd7
2 of 14
No
No
Rnor\Adgrb1
1 of 14
No
No
Rnor\Adgrb2
1 of 14
No
No
Rnor\Adgrl3
1 of 14
No
No
Rnor\Fzd10
1 of 14
No
Yes
Rnor\Mfrp
1 of 14
No
No
Rnor\Sfrp1
1 of 14
No
No
Rnor\Sfrp2
1 of 14
No
No
Rnor\Sfrp5
1 of 14
No
No
Mus musculus (laboratory mouse) (21)
Mmus\Smo
13 of 14
Yes
Yes
Mmus\Fzd4
3 of 14
No
No
Mmus\Fzd8
3 of 14
No
No
Mmus\Fzd9
3 of 14
No
No
Mmus\Fzd10
3 of 14
No
No
Mmus\Frzb
2 of 14
No
No
Mmus\Fzd1
2 of 14
No
No
Mmus\Fzd2
2 of 14
No
No
Mmus\Fzd3
2 of 14
No
No
Mmus\Fzd5
2 of 14
No
No
Mmus\Fzd6
2 of 14
No
No
Mmus\Fzd7
2 of 14
No
No
Mmus\Adgrb1
1 of 14
No
No
Mmus\Adgrb2
1 of 14
No
No
Mmus\Adgrl3
1 of 14
No
No
Mmus\Corin
1 of 14
No
No
Mmus\Mfrp
1 of 14
No
No
Mmus\Sfrp1
1 of 14
No
No
1  
Mmus\Sfrp2
1 of 14
No
No
Mmus\Sfrp4
1 of 14
No
No
Mmus\Sfrp5
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (16)
Xtro\smo
8 of 13
Yes
Yes
Xtro\fzd4
2 of 13
No
No
Xtro\fzd5
2 of 13
No
No
Xtro\fzd9
2 of 13
No
No
Xtro\cradd.1
1 of 13
No
Yes
Xtro\frzb
1 of 13
No
No
Xtro\fzd3
1 of 13
No
No
Xtro\fzd6
1 of 13
No
No
Xtro\fzd7
1 of 13
No
No
Xtro\fzd8
1 of 13
No
No
Xtro\fzd10
1 of 13
No
No
Xtro\galnt6.3
1 of 13
No
Yes
Xtro\hsdl2
1 of 13
No
No
Xtro\pik3r3
1 of 13
No
No
Xtro\stoml2
1 of 13
No
No
Xtro\tlx1
1 of 13
No
No
Danio rerio (Zebrafish) (24)
Drer\smo
13 of 14
Yes
Yes
Drer\fzd5
3 of 14
No
No
Drer\fzd8a
3 of 14
No
No
Drer\fzd8b
3 of 14
No
No
Drer\fzd9b
3 of 14
No
No
Drer\fzd10
3 of 14
No
No
Drer\frzb
2 of 14
No
No
Drer\fzd2
2 of 14
No
No
Drer\fzd3a
2 of 14
No
No
Drer\fzd3b
2 of 14
No
No
Drer\fzd4
2 of 14
No
No
Drer\fzd7a
2 of 14
No
No
Drer\fzd7b
2 of 14
No
No
Drer\fzd9a
2 of 14
No
No
Drer\adgrb3
1 of 14
No
Yes
Drer\adgrl4
1 of 14
No
No
Drer\col18a1a
1 of 14
No
No
Drer\fzd1
1 of 14
No
No
Drer\fzd6
1 of 14
No
No
Drer\mfrp
1 of 14
No
No
Drer\sfrp1a
1 of 14
No
No
Drer\sfrp1b
1 of 14
No
No
Drer\sfrp2
1 of 14
No
No
Drer\sfrp5
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (5)
Cele\cfz-2
2 of 14
Yes
No
Cele\mom-5
2 of 14
Yes
No
Cele\lin-17
1 of 14
No
No
Cele\mig-1
1 of 14
No
No
Cele\sfrp-1
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (9)
Agam\GPRSMO
11 of 12
Yes
Yes
Agam\GPRFZ2
3 of 12
No
No
Agam\GPRFZ3
3 of 12
No
No
Agam\GPRFZ4
3 of 12
No
No
Agam\AgaP_AGAP000303
1 of 12
No
No
Agam\AgaP_AGAP011379
1 of 12
No
No
Agam\GPRFZ1A
1 of 12
No
Yes
Agam\GPRFZ1B
1 of 12
No
Yes
Agam\SCRASP2
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:smo. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (7)
fz2
5 of 13
fz3
5 of 13
fz4
5 of 13
fz
4 of 13
CG1632
1 of 13
Corin
1 of 13
mthl15
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Allele
Disease
Evidence
References
smoRNAi.UAS.cUa
model of  primary bacterial infectious disease
CEA
(Lee et al., 2015)
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 1 )
Allele
Disease
Interaction
References
smo1
ameliorates  neurodegenerative disease
modeled by ptc559.1
(Gazi et al., 2009)
Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
Homo sapiens (Human)
Gene name
Score
OMIM
OMIM Phenotype
DO term
Complementation?
Transgene?
SMO; smoothened, frizzled class receptor
13 of 14
601500
    FZD4; frizzled class receptor 4
    3 of 14
    604579
    FZD5; frizzled class receptor 5
    3 of 14
    601723
      FZD8; frizzled class receptor 8
      3 of 14
      606146
          FZD9; frizzled class receptor 9
          3 of 14
          601766
              FZD10; frizzled class receptor 10
              3 of 14
              606147
                  Functional Complementation Data
                  Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
                  Interactions
                  Summary of Physical Interactions
                  Interaction Browsers
                  View in FlyBase Interactions Browser   View in MIST tool (external link)
                  Please see the Physical Interaction reports below for full details
                  protein-protein
                  Physical Interaction
                  Assay
                  References
                  smo
                  pull down, anti tag western blot, fluorescent resonance energy transfer, anti tag coimmunoprecipitation, fluorescence microscopy, anti bait coimmunoprecipitation, western blot, autoradiography, bioluminescence resonance energy transfer, fluorescence technology
                  (Jiang et al., 2016, Maier et al., 2014, Fan et al., 2012, Chen et al., 2010, Walthall et al., 2007, Zhao et al., 2007)
                  smo - Cul4
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Li et al., 2018)
                  smo - Gprk2
                  anti tag coimmunoprecipitation, anti tag western blot, fluorescent resonance energy transfer, fluorescence microscopy, pull down
                  (Jiang et al., 2016, Chen et al., 2010)
                  smo - Gβ13F
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Li et al., 2018)
                  smo - Herc4
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Jiang et al., 2019)
                  smo - Hrs
                  pull down, anti tag western blot, anti tag coimmunoprecipitation, western blot
                  (Zhou et al., 2018, Fan et al., 2013)
                  smo - Pka-C1
                  anti tag coimmunoprecipitation, anti tag western blot, bimolecular fluorescence complementation, fluorescence microscopy, western blot, anti bait coimmunoprecipitation
                  (Fan et al., 2019, Chen et al., 2017, Li et al., 2014, Ranieri et al., 2014)
                  smo - Pp1-13C
                  anti tag coimmunoprecipitation, western blot
                  (Su et al., 2011)
                  smo - Pp1-87B
                  anti tag coimmunoprecipitation, western blot
                  (Su et al., 2011)
                  smo - Pp1α-96A
                  anti tag coimmunoprecipitation, western blot
                  (Su et al., 2011)
                  smo - Pp4-19C
                  anti tag coimmunoprecipitation, anti tag western blot, pull down
                  (Jia et al., 2009)
                  smo - Rack1
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Li et al., 2020)
                  smo - Smurf
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Li et al., 2018, Huang et al., 2013)
                  smo - Sumo
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Zhang et al., 2017)
                  smo - Sxl
                  anti bait coimmunoprecipitation, western blot
                  (Walthall et al., 2007)
                  smo - Ubi-p5E
                  anti tag coimmunoprecipitation, covalent binding, anti tag western blot
                  (Yang et al., 2013)
                  smo - Ubi-p63E
                  anti tag coimmunoprecipitation, covalent binding, anti tag western blot
                  (Yang et al., 2013)
                  smo - Uch-L5
                  anti bait coimmunoprecipitation, anti tag western blot, anti tag coimmunoprecipitation
                  (Zhou et al., 2018)
                  smo - Uch-L5R
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Zhou et al., 2018)
                  smo - Ulp1
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Zhang et al., 2017, Ma et al., 2016)
                  smo - Usp8
                  anti tag coimmunoprecipitation, anti tag western blot, pull down
                  (Li et al., 2020, Ma et al., 2016, Fan et al., 2013, Li et al., 2012, Xia et al., 2012)
                  smo - Vps36
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Yang et al., 2013)
                  smo - ci
                  anti bait coimmunoprecipitation, peptide massfingerprinting, western blot
                  (Walthall et al., 2007, Lum et al., 2003, Ruel et al., 2003)
                  smo - cos
                  bioluminescence resonance energy transfer, fluorescence technology, anti tag coimmunoprecipitation, western blot, anti bait coimmunoprecipitation, anti tag western blot, colocalization, fluorescence microscopy, inferred by author, tag visualisation, fluorescent resonance energy transfer, pull down, two hybrid, peptide massfingerprinting
                  (Li et al., 2014, Maier et al., 2014, Ranieri et al., 2014, Fan et al., 2012, Ranieri et al., 2012, Shi et al., 2011, Zhang et al., 2011, Liu et al., 2007, Ruel et al., 2007, Walthall et al., 2007, Zhang et al., 2005, Jia et al., 2003, Lum et al., 2003, Ogden et al., 2003, Ruel et al., 2003)
                  smo - flw
                  anti tag coimmunoprecipitation, western blot
                  (Su et al., 2011)
                  smo - fu
                  anti bait coimmunoprecipitation, western blot, bimolecular fluorescence complementation, fluorescence microscopy, two hybrid, anti tag coimmunoprecipitation, anti tag western blot, colocalization, inferred by author, peptide massfingerprinting
                  (Kubíková et al., 2023, Giordano et al., 2018, Ranieri et al., 2014, Ranieri et al., 2012, Malpel et al., 2007, Jia et al., 2003, Lum et al., 2003, Ogden et al., 2003, Ruel et al., 2003)
                  smo - gish
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Li et al., 2016)
                  smo - krz
                  anti tag coimmunoprecipitation, western blot, anti tag western blot
                  (Jiang et al., 2018, Zhang et al., 2017, Li et al., 2012, Molnar et al., 2011)
                  smo - mts
                  anti tag coimmunoprecipitation, western blot
                  (Su et al., 2011)
                  smo - pic
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Li et al., 2018)
                  smo - ptc
                  anti tag coimmunoprecipitation, western blot, anti bait coimmunoprecipitation
                  (Walthall et al., 2007, Johnson et al., 2000)
                  smo - smg
                  colocalization, fluorescence microscopy, inferred by author, anti tag coimmunoprecipitation, anti tag western blot, pull down, molecular weight estimation by staining, isothermal titration calorimetry, predetermined participant, x-ray crystallography, two hybrid
                  (Kubíková et al., 2023, Bruzzone et al., 2020)
                  smo - wdb
                  pull down, western blot, anti tag coimmunoprecipitation
                  (Su et al., 2011)
                  RNA-RNA
                  Physical Interaction
                  Assay
                  References
                  smo - mir-12
                  fluorescence technology
                  (Friggi-Grelin et al., 2008)
                  smo - mir-283
                  fluorescence technology
                  (Friggi-Grelin et al., 2008)
                  smo - mir-5
                  immunohistochemistry, luminiscence technology, necessary binding region, fluorescence technology
                  (Wu et al., 2012)
                  smo - mir-960
                  immunocytochemistry, luminiscence technology, necessary binding region, fluorescence technology
                  (Gao et al., 2013)
                  Summary of Genetic Interactions
                  Interaction Browsers
                  View in FlyBase Interactions Browser   View in MIST tool (external link)
                  Please look at the allele data for full details of the genetic interactions
                  Starting gene(s)
                  Interaction type
                  Interacting gene(s)
                  Reference
                  smo
                  enhanceable
                  apt
                  (Wang et al., 2017)
                  smo
                  suppressible
                  Cad99C
                  (Lee et al., 2015)
                  smo
                  suppressible
                  ci
                  (Fu and Baker, 2003)
                  smo
                  enhanceable
                  CkIIα
                  (Jia et al., 2010)
                  smo
                  suppressible
                  cos
                  (Hooper, 2003)
                  smo
                  enhanceable
                  cos
                  (Ogden et al., 2006)
                  smo
                  suppressible
                  dlp
                  (Tu et al., 2020)
                  smo
                  suppressible
                  dpp|eya
                  (Pappu et al., 2003)
                  smo
                  enhanceable
                  dsh
                  (Dai et al., 2017)
                  smo
                  enhanceable
                  E(smoDN)111
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)198
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)203
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)219
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)390
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)391
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)418
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)461
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)474
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)529
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)548
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)786
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)830
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)845
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)871
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)918
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)986
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1124
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1131
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1171
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1333
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1338
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1411
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1414
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1419
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1422
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1438
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1485
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1496
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1551
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1575
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1644
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1700
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1736
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1823
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1824
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1835
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1857
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1875
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)1889
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2016
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2038
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2042
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2047
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2070
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2101
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2117
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2135
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2162
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2169
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2243
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2281
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2364
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2515
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2546
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2576
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2577
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2578
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2583
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2590
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2619
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)2640
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)A
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  enhanceable
                  E(smoDN)B-right
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  enhanceable
                  E(smoDN)E
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  enhanceable
                  E(smoDN)G
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  enhanceable
                  E(smoDN)M
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  enhanceable
                  E(smoDN)N
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  E(smoDN)O
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  eIF1A
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  enhanceable
                  eRF1
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  enhanceable
                  fu
                  (Hooper, 2003)
                  smo
                  suppressible
                  fu
                  (Malpel et al., 2007, Hooper, 2003)
                  smo
                  suppressible
                  gig
                  (Kim et al., 2017)
                  smo
                  enhanceable
                  gish
                  (Li et al., 2016)
                  smo
                  suppressible
                  gish
                  (Li et al., 2016)
                  smo
                  enhanceable
                  Gprk2
                  (Li et al., 2016, Chen et al., 2010)
                  smo|gish
                  enhanceable
                  Gprk2
                  (Li et al., 2016)
                  smo
                  enhanceable
                  hh
                  (Wang et al., 2017, Casso et al., 2008)
                  smo
                  suppressible
                  Hrs
                  (Fan et al., 2013)
                  smo
                  enhanceable
                  Hrs
                  (Fan et al., 2013)
                  smo
                  enhanceable
                  jaft
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  suppressible
                  Kdm3
                  (Shalaby et al., 2017)
                  smo
                  enhanceable
                  Kdm4A
                  (Shalaby et al., 2017)
                  smo
                  enhanceable
                  Kdm4B
                  (Shalaby et al., 2017)
                  smo
                  enhanceable
                  krz
                  (Molnar et al., 2011)
                  smo
                  enhanceable
                  lwr
                  (Zhang et al., 2017)
                  smo
                  enhanceable
                  M(smoDN)K
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  suppressible
                  M(smoDN)K
                  (Collins, 2005.7.7)
                  smo
                  enhanceable
                  Mad
                  (Fu and Baker, 2003)
                  smo
                  enhanceable
                  mirr
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  enhanceable
                  mts
                  (Casso et al., 2008)
                  smo
                  enhanceable
                  pav
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  suppressible
                  pav
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  suppressible
                  ptc
                  (Casso et al., 2008, Collins and Cohen, 2005, Hooper, 2003)
                  smo
                  enhanceable
                  ptc
                  (Collins, 2005.7.7)
                  smo
                  suppressible
                  S(smoDN)166
                  (Collins, 2005.7.7)
                  smo
                  suppressible
                  S(smoDN)189
                  (Collins, 2005.7.7)
                  smo
                  suppressible
                  S(smoDN)194
                  (Collins, 2005.7.7)
                  smo
                  suppressible
                  S(smoDN)204
                  (Collins, 2005.7.7)
                  smo
                  suppressible
                  S(smoDN)395
                  (Collins, 2005.7.7)
                  smo
                  suppressible
                  S(smoDN)1378
                  (Collins, 2005.7.7)
                  smo
                  suppressible
                  S(smoDN)1657
                  (Collins, 2005.7.7)
                  smo
                  suppressible
                  S(smoDN)2472
                  (Collins, 2005.7.7)
                  smo
                  suppressible
                  S(smoDN)2629
                  (Collins, 2005.7.7)
                  Pka-C1|smo
                  suppressible
                  sgg
                  (Price and Kalderon, 2002)
                  smo
                  suppressible
                  so|eya
                  (Pappu et al., 2003)
                  smo
                  suppressible
                  Stat92E
                  (Liu et al., 2015)
                  smo
                  enhanceable
                  Su(var)2-10
                  (Zhang et al., 2017)
                  smo
                  enhanceable
                  Sumo
                  (Zhang et al., 2017)
                  smo
                  suppressible
                  swm
                  (Casso et al., 2008)
                  smo
                  enhanceable
                  tkv
                  (Fu and Baker, 2003)
                  smo
                  enhanceable
                  Trs20
                  (Collins, 2005.7.7, Collins and Cohen, 2005)
                  smo
                  enhanceable
                  TSG101
                  (Fan et al., 2013)
                  smo
                  suppressible
                  Ulp1
                  (Zhang et al., 2017)
                  smo
                  suppressible
                  wg
                  (Quirk et al., 1997)
                  Starting gene(s)
                  Interaction type
                  Interacting gene(s)
                  Reference
                  boi
                  suppressible
                  smo
                  (Hartman et al., 2010)
                  cos
                  enhanceable
                  smo
                  (Ogden et al., 2006)
                  Delta
                  enhanceable
                  smo
                  (Da Ros et al., 2013)
                  dsh
                  enhanceable
                  smo
                  (Dai et al., 2017)
                  E2f1
                  suppressible
                  smo
                  (Ambrus et al., 2007)
                  fu
                  suppressible
                  smo
                  (Claret et al., 2007)
                  Gprk2
                  enhanceable
                  smo
                  (Molnar et al., 2007)
                  hid
                  enhanceable
                  smo
                  (Christiansen et al., 2013)
                  krz
                  suppressible
                  smo
                  (Molnar et al., 2011)
                  L
                  suppressible
                  smo
                  (Singh et al., 2005)
                  Moe
                  enhanceable
                  smo
                  (Molnar and de Celis, 2006)
                  pnr
                  enhanceable
                  smo
                  (Pena-Rangel et al., 2002)
                  ptc
                  suppressible
                  smo
                  (Gazi et al., 2009, Merianda et al., 2005, Mullor and Guerrero, 2000, Quirk et al., 1997)
                  ptc
                  enhanceable
                  smo
                  (Hooper, 2013.5.30, Shyamala and Bhat, 2002)
                  slmb
                  suppressible
                  smo
                  (Theodosiou et al., 1998)
                  TkR99D
                  suppressible
                  smo
                  (Im et al., 2015)
                  External Data
                  Subunit Structure (UniProtKB)
                  Interacts with cos.
                  (UniProt, P91682 )
                  Linkouts
                  BioGRID - A database of protein and genetic interactions.
                  DroID - A comprehensive database of gene and protein interactions.
                  MIST (genetic) - An integrated Molecular Interaction Database
                  MIST (protein-protein) - An integrated Molecular Interaction Database
                  Pathways
                  Signaling Pathways (FlyBase)
                  Metabolic Pathways
                  FlyBase
                  External Links
                  External Data
                  Linkouts
                  KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
                  Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                  SignaLink - A signaling pathway resource with multi-layered regulatory networks.
                  Class of Gene
                  Genomic Location and Detailed Mapping Data
                  Chromosome (arm)
                  2L
                  Recombination map
                  2-0.5
                  Cytogenetic map
                  21B7-21B7
                  Sequence location
                  FlyBase Computed Cytological Location
                  Cytogenetic map
                  Evidence for location
                  21B7-21B7
                  Limits computationally determined from genome sequence between P{lacW}spenk06805&P{lacW}l(2)k13604k13604 and P{lacW}U2af38k14504
                  Experimentally Determined Cytological Location
                  Cytogenetic map
                  Notes
                  References
                  Experimentally Determined Recombination Data
                  Location

                  2-0.5

                  (FlyBase, 2016)

                  2-0.0

                  (Comeron et al., 2012)

                  2-4

                  (Tearle and Nusslein-Volhard, 1987)

                  2-0.4

                  Left of (cM)
                  Right of (cM)
                  Notes
                  Stocks and Reagents
                  Stocks (41)
                  Genomic Clones (18)
                   

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

                  cDNA Clones (36)
                   

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

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

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

                  cDNA Clones, End Sequenced (ESTs)
                  BDGP DGC clones
                  Other clones
                  RNAi and Array Information
                  Linkouts
                  DRSC - Results frm RNAi screens
                  Antibody Information
                  Laboratory Generated Antibodies
                   

                  polyclonal

                  (Li et al., 2016, Maier et al., 2014, Ranieri et al., 2014, Fan et al., 2012, Xia et al., 2012, Su et al., 2011)
                  Commercially Available Antibodies
                   
                  Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
                  Cell Line Information
                  Publicly Available Cell Lines
                   
                    Other Stable Cell Lines
                     

                    • New stable cell line derived from S2-unspecified : A stable S2 cell line was isolated that expresses the smoIP fluorescence-based sensor under the control of UAS and reproducibly responds to hh stimulation.

                      (Albert and Bökel, 2017)

                    • New stable cell line derived from S2-unspecified : S2 cells stably expressing an inducible Tag:MYC-tagged smo transgene under the control of the metallothionein promoter (pMT-Myc-Smo) were created. S2 cells stably expressing the N-terminal fragment of hhh protein (HhN) that is active for signaling were created.

                      (Li et al., 2018)
                    Other Comments

                    smo protein promotes the relocalisation of fu protein from vesicles to the plasma membrane in response to hh signaling and reciprocally, fu protein controls the stability and phosphorylation of smo protein.

                    (Claret et al., 2007)

                    A region in the C-terminus of smo protein is sufficient for its interaction with fu protein (spans amino acids 985 to 1036).

                    (Malpel et al., 2007)

                    Phosphorylation activates smo protein by inducing a conformational switch. This occurs by antagonising multiple Arg clusters in the smo cytoplasmic tail. The Arg clusters inhibit smo protein by blocking its cell surface expression and keeping it in an inactive conformation that is maintained by electrostatic interactions. Phosphorylation disrupts the interactions and induces a conformational switch and dimerisation of smo protein cytoplasmic tails, which is essential for pathway activation.

                    (Zhao et al., 2007)

                    Identified as a component of the hh signalling pathway in a genome-wide RNAi screen. dsRNA made from templates generated with primers directed affects the extent of expression of a hh signaling reporter construct in Clone 8 cells.

                    (Nybakken et al., 2005)

                    smo protein accumulates in the plasma membrane of cells that lack ptc activity, whereas when co-expressed with ptc, smo protein localises exclusively intracellularly, in distinct punctate structures.

                    (Nakano et al., 2004)

                    dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.

                    (Kiger et al., 2003)

                    dsRNA made from templates generated with primers against this gene tested in a cl-8 cell reporter assay to examine any role of smo in the hh signalling pathway.

                    (Lum, 2003)

                    The last 301 amino acids of the cos COOH terminus are sufficient to provide full association with smo.

                    (Ruel et al., 2003)

                    Epistatic analysis places cos function downstream of ptc and smo.

                    (Ruel et al., 2003)

                    dsRNA corresponding to smo used in S2 cultured cell assay.

                    (Ruel et al., 2003)

                    17 alleles of smo have been recovered in a screen for mutations with smo-like mutant phenotypes in clones in the wing.

                    (Vegh and Basler, 2003)

                    Control of smo localisation appears to be a crucial step in hh signalling.

                    (Zhu et al., 2003)

                    ptc-smo signalling has a role in head morphogenesis to promote cell proliferation via activation of babo.

                    (Shyamala and Bhat, 2002)

                    ptc protein destabilises smo protein in the absence of hh protein.

                    (Denef et al., 2000)

                    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.

                    (Therond et al., 1999)

                    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.

                    (Chen and Struhl, 1998)

                    Anterior cells in the wing that lack smo function no longer obey a lineage restriction in the normal position of the anterior-posterior boundary.

                    (Blair and Ralston, 1997)

                    Complementation and epistasis tests indicate that oro is not allelic to smo.

                    (Epps et al., 1997)

                    smo encodes a serpentine protein highly conserved in evolution.

                    (Quirk et al., 1997)

                    smo activity is required for transduction of hh but not wg. smo acts downstream from ptc to transduce the hh signal.

                    (Quirk et al., 1997)

                    Loss of smo function causes a hh-like phenotype.

                    (Quirk et al., 1997)

                    Morphogenetic furrow progression is significantly delayed, but not prevented, in smo clones.

                    (Strutt and Mlodzik, 1997)

                    smo is a putative receptor of the hh signal.

                    (Alcedo et al., 1996)

                    smo encodes a seven-pass membrane protein. In direction of increasing cytology: anon-21Ca+ anon-21Cb- smo+ anon-21Cc-

                    (Alcedo et al., 1996)

                    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.

                    (Chen and Struhl, 1996)

                    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.

                    (Stone et al., 1996)

                    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.

                    (van den Heuvel and Ingham, 1996)

                    The effects of removing smo gene activity from embryos in which hh or wg is ectopically expressed suggest a role for smo in the hh signalling pathway.

                    (van den Heuvel and Ingham, 1995)

                    All alleles are zygotic lethal, cold sensitive, at the lower temperature the phenotype of all alleles is quite variable suggesting that the gene may be maternally expressed.

                    (van den Heuvel and Ingham, 1995)

                    Direct wg autoregulation differs from wg signalling to adjacent cells in the importance of fu, smo and ci relative to sgg and arm.

                    (Hooper, 1994)

                    The smo gene acts downstream of wg in pattern formation.

                    (Sampedro et al., 1993)

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

                    (Parkhurst and Ish-Horowicz, 1991)
                    Relationship to Other Genes
                    Source for database merge of
                    Additional comments
                    Nomenclature History
                    Source for database identify of
                    Nomenclature comments
                    Etymology

                    Named "smoothened" for the smooth, denticle-free cuticle in mutant embryos.

                    (Nusslein-Volhard and Wieschaus, 1980)
                    Synonyms and Secondary IDs (9)
                    Reported As
                    Symbol Synonym
                    CG11561
                    (National Institute of Genetics Fly Stocks, 2022-, Meadows, 2017.1.11, Hosono et al., 2015, Ni et al., 2010.12.1, Perkins et al., 2009, Perkins et al., 2009.3.31, Nybakken et al., 2005, Kim et al., 2004)
                    SMO
                    (Zhang et al., 2021, Percival-Smith et al., 2017, Sanial et al., 2017, Brigui et al., 2015, Marques-Pita and Rocha, 2013, Tran et al., 2013, Dahmann et al., 2011, Ingham et al., 2011, Stoll et al., 2010, Subramanian and Gadgil, 2010, Malpel et al., 2007, Plessis et al., 2007, Sanial and Plessis, 2007, Zhao et al., 2007)
                    Smo
                    (Goins et al., 2024, Jang et al., 2024, Chang, 2022, Han and Jiang, 2022, Emmons-Bell and Hariharan, 2021, Liu et al., 2021, Manikowski et al., 2021, Saad and Hipfner, 2021, Surkova et al., 2021, Zhang and Jiang, 2021, Zhao et al., 2021, González-Méndez et al., 2020, Gorla and Bashaw, 2020, Gou et al., 2020, Kamdem et al., 2020, Lan et al., 2020, Li et al., 2020, Luo et al., 2020, Nishihara, 2020, Pan et al., 2020, Prince et al., 2020, Zhang et al., 2020, Fan et al., 2019, Han et al., 2019, Jiang et al., 2019, Meltzer, 2019-, Meltzer et al., 2019, Giordano et al., 2018, Jiang et al., 2018, Lehmann, 2018, Li et al., 2018, Li et al., 2018, Zhou et al., 2018, Chen et al., 2017, Hsia et al., 2017, Li et al., 2017, Shalaby et al., 2017, Zhang et al., 2017, Zhao et al., 2017, Jiang et al., 2016, Lee et al., 2016, Li et al., 2016, Mbodj et al., 2016, Ulmschneider et al., 2016, Dorn and Dorn, 2015, Gurdziel et al., 2015, Khaliullina et al., 2015, Lee et al., 2015, Marada et al., 2015, Oh et al., 2015, Sharpe et al., 2015, Xiong et al., 2015, Jiang et al., 2014, Kuzhandaivel et al., 2014, Pichaud, 2014, Shi et al., 2014, Wang et al., 2014, Zhang et al., 2014, Bausek, 2013, Briscoe and Thérond, 2013, Chai et al., 2013, Chen and Jiang, 2013, Gao et al., 2013, Gao et al., 2013, Mbodj et al., 2013, Palm et al., 2013, Shi et al., 2013, Yang et al., 2013, Yousefian et al., 2013, Zhang et al., 2013, Cheng et al., 2012, Fan et al., 2012, Legent et al., 2012, Li et al., 2012, Robbins et al., 2012, Rojas-Ríos et al., 2012, Xia et al., 2012, Shi et al., 2011, Zhang et al., 2011, Chen et al., 2010, Jia et al., 2010, Mukai et al., 2010, Zheng et al., 2010, Ayers et al., 2009, Farzan et al., 2009, Mao and Freeman, 2009, González et al., 2008, Liu et al., 2008, Su et al., 2008, Wu and Mlodzik, 2008, Zhao and Jiang, 2008, Liu et al., 2007, Molnar et al., 2007, Walthall et al., 2007, Zhao et al., 2007, Beenken and Mohammadi, 2006, Fisher and Howie, 2006, Jiang, 2006, Ogden et al., 2006, Osterlund and Kogerman, 2006, Wilson and Chuang, 2006, Dawber et al., 2005, Jekely and Rorth, 2003, Kuwabara and Labouesse, 2002, Chen et al., 1999)
                    dSmo
                    (Sun et al., 2019, Chen and Jiang, 2013, Chen et al., 2009, De Rivoyre et al., 2006, Stone et al., 1996)
                    smo
                    (Port et al., 2026, Bischoff et al., 2025, Deshpande et al., 2025, Kong et al., 2025, Deshpande et al., 2023, Bolatto et al., 2022, Bostock et al., 2022, Chen et al., 2022, Gonçalves Antunes et al., 2022, He et al., 2022, Míguez et al., 2022, National Institute of Genetics Fly Stocks, 2022-, Schlichting et al., 2022, Yang et al., 2022, Boukhatmi, 2021, Everetts et al., 2021, Liu et al., 2021, Liu et al., 2021, Tatapudy et al., 2021, Tu et al., 2021, Basu et al., 2020, Bruzzone et al., 2020, Little et al., 2020, Miller et al., 2020, Port et al., 2020, Tu et al., 2020, Wang and Dahmann, 2020, Brodskiy et al., 2019, Li and Baker, 2019, Xu et al., 2019, Ahaley, 2018, Baldeosingh et al., 2018, Billmann et al., 2018, Tseng et al., 2018, Albert and Bökel, 2017, Dai et al., 2017, Garcia-Garcia et al., 2017, González-Méndez et al., 2017, Lai et al., 2017, Lu et al., 2017, Recasens-Alvarez et al., 2017, Sanial et al., 2017, Solis et al., 2017, Transgenic RNAi Project members, 2017-, Li et al., 2016, Ma et al., 2016, Sarov et al., 2016, Han et al., 2015, Im et al., 2015, Li et al., 2015, Liu et al., 2015, Lu et al., 2015, Matsuda et al., 2015, Rudolf et al., 2015, Sîrbu et al., 2015, Zhou et al., 2015, Blaquiere et al., 2014, Camp et al., 2014, Huang and Kalderon, 2014, Issman-Zecharya and Schuldiner, 2014, Karandikar et al., 2014, Li et al., 2014, Li et al., 2014, Liu et al., 2014, Maier et al., 2014, Piñeiro et al., 2014, Ranieri et al., 2014, Atkins et al., 2013, Avanesov and Blair, 2013, Bilioni et al., 2013, Chai et al., 2013, Da Ros et al., 2013, Fan et al., 2013, Geisbrecht et al., 2013, Hartman et al., 2013, Hooper, 2013.5.30, Hooper, 2013.6.26, Huang et al., 2013, Kupinski et al., 2013, Li et al., 2013, Nedelcu et al., 2013, Rana et al., 2013, Saunders et al., 2013, Spratford and Kumar, 2013, Yu et al., 2013, Zhang et al., 2013, Carroll et al., 2012, Cheng et al., 2012, Christiansen et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Jimenez-Sanchez et al., 2012, Krzemien et al., 2012, Rojas-Ríos et al., 2012, Wang et al., 2012, Wu et al., 2012, Bhattacharya and Baker, 2011, Marks and Kalderon, 2011, Molnar et al., 2011, Nicholson et al., 2011, Schilling et al., 2011, Su et al., 2011, Terriente-Félix et al., 2011, Toku et al., 2011, Zhang et al., 2011, Zhou and Kalderon, 2011, Biehs et al., 2010, Camp et al., 2010, Cheng et al., 2010, Chou et al., 2010, de Celis and Molnar, 2010, Firth et al., 2010, Hartman et al., 2010, Jia et al., 2010, Li et al., 2010, Lopes and Casares, 2010, Pospisilik et al., 2010, Raisin et al., 2010, Terriente-Félix et al., 2010, Wasbrough et al., 2010, Yavari et al., 2010, Zhou and Kalderon, 2010, Baker et al., 2009, Baron et al., 2009, Blanco et al., 2009, Deshpande et al., 2009, Firth and Baker, 2009, Foronda et al., 2009, Gazi et al., 2009, Jia et al., 2009, Khaliullina et al., 2009, Landsberg et al., 2009, Mosimann et al., 2009, Renault et al., 2009, Vied and Kalderon, 2009, Casso et al., 2008, Chen et al., 2008, Fan and Bergmann, 2008, Farzan et al., 2008, Friggi-Grelin et al., 2008, Gallet et al., 2008, Lim et al., 2008, Melicharek et al., 2008, Ogden et al., 2008, Schlichting and Dahmann, 2008, Tanaka-Matakatsu and Du, 2008, Terriente Felix and de Celis, 2008.1.16, Wang and Price, 2008, Ambrus et al., 2007, Bashaw, 2007, Bejarano et al., 2007, Beltran et al., 2007, Casso et al., 2007, Chen et al., 2007, Claret et al., 2007, Cook, 2007.7.24, Corrigall et al., 2007, Escudero and Freeman, 2007, Escudero et al., 2007, Goodfellow et al., 2007, Lecuyer et al., 2007, Molnar et al., 2007, Ruel et al., 2007, Smelkinson et al., 2007, Sweeney et al., 2007, Colosimo and Tolwinski, 2006, Croker et al., 2006, D'Costa et al., 2006, Kent et al., 2006, Lu et al., 2006, McLellan et al., 2006, Molnar and de Celis, 2006, Sisson et al., 2006, Smelkinson and Kalderon, 2006, Suh et al., 2006, Umetsu et al., 2006, Varjosalo et al., 2006, Vrailas and Moses, 2006, Vrailas et al., 2006, Yao et al., 2006, Zhou et al., 2006, Baonza and Freeman, 2005, Briscoe and Therond, 2005, Cabernard and Affolter, 2005, Deshpande and Schedl, 2005, Firth and Baker, 2005, Firth and Baker, 2005, Ishii, 2005, Jia et al., 2005, Kirilly et al., 2005, Merianda et al., 2005, Nybakken et al., 2005, Torroja et al., 2005, Wei et al., 2005, Xie et al., 2005, Zhang et al., 2005, Punzo et al., 2004, Raabe et al., 2004)
                    Name Synonyms
                    Smoothene
                    (Hartman et al., 2010)
                    Smoothened
                    (Harnish et al., 2021, Jafari and Alenius, 2021, Cui et al., 2020, Nishihara, 2020, Palm and Rodenfels, 2020, Xu et al., 2019, Zhao et al., 2017, Ulmschneider et al., 2016, Lee et al., 2015, Wang et al., 2014, Bausek, 2013, Chen and Jiang, 2013, Fan et al., 2013, Huang et al., 2013, Shi et al., 2013, Wong et al., 2013, Yang et al., 2013, Cheng et al., 2012, Fan et al., 2012, Li et al., 2012, Rojas-Ríos et al., 2012, Molnar et al., 2011, Schilling et al., 2011, Shi et al., 2011, Camp et al., 2010, Chou et al., 2010, Jia et al., 2010, Mukai et al., 2010, Yavari et al., 2010, Zhou and Kalderon, 2010, Ayers et al., 2009, Blanco et al., 2009, Farzan et al., 2009, Gazi et al., 2009, Jia et al., 2009, Khaliullina et al., 2009, Mosimann et al., 2009, Farzan et al., 2008, Liu et al., 2008, Su et al., 2008, Wu and Mlodzik, 2008, Bangi et al., 2007, Bejarano et al., 2007, Casso et al., 2007, Claret et al., 2007, Escudero and Freeman, 2007, Malpel et al., 2007, Molnar et al., 2007, Plessis et al., 2007, Sanial and Plessis, 2007, Walthall et al., 2007, Zhao et al., 2007, Beenken and Mohammadi, 2006, Colosimo and Tolwinski, 2006, Maitra et al., 2006, Ogden et al., 2006, Panakova and Eaton, 2006, Wilson and Chuang, 2006, Dawber et al., 2005, Torroja et al., 2005)
                    smooth
                    (Sampedro et al., 1993, Parkhurst and Ish-Horowicz, 1991, Tearle and Nusslein-Volhard, 1987)
                    smoothened
                    (Li et al., 2017, Shalaby et al., 2017, Wieschaus and Nüsslein-Volhard, 2016, Matsuda et al., 2015, Xiong et al., 2015, Piñeiro et al., 2014, Ranieri et al., 2014, Avanesov and Blair, 2013, Da Ros et al., 2013, Nedelcu et al., 2013, Rana et al., 2013, Spratford and Kumar, 2013, Krzemien et al., 2012, Bhattacharya and Baker, 2011, Marks and Kalderon, 2011, Nicholson et al., 2011, Zhou and Kalderon, 2011, Cheng et al., 2010, de Celis and Molnar, 2010, Lopes and Casares, 2010, Terriente-Félix et al., 2010, Baron et al., 2009, Foronda et al., 2009, Landsberg et al., 2009, Casso et al., 2008, Fan and Bergmann, 2008, Lim et al., 2008, Ogden et al., 2008, Corrigall et al., 2007, Escudero et al., 2007, Sweeney et al., 2007, Zhao et al., 2007, Jones et al., 2006, Vrailas and Moses, 2006, Yao et al., 2006, Baonza and Freeman, 2005, Merianda et al., 2005, Hime et al., 2004, Hooper, 1997.1.29)
                    Secondary FlyBase IDs
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                      External Crossreferences and Linkouts ( 54 )
                      Sequence Crossreferences
                      NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
                      GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
                      GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
                      RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
                      UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
                      UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
                      UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
                      Other crossreferences
                      AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
                      BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
                      DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
                      EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
                      FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
                      FlyMine - An integrated database for Drosophila genomics
                      InterPro - A database of protein families, domains and functional sites
                      KEGG Genes - Molecular building blocks of life in the genomic space.
                      MARRVEL_MODEL - MARRVEL (model organism gene)
                      PDB - An information portal to biological macromolecular structures
                      Linkouts
                      BioGRID - A database of protein and genetic interactions.
                      Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
                      DroID - A comprehensive database of gene and protein interactions.
                      DRSC - Results frm RNAi screens
                      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
                      Flygut - An atlas of the Drosophila adult midgut
                      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
                      Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                      SignaLink - A signaling pathway resource with multi-layered regulatory networks.
                      References (719)