FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\Smox
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General Information
Symbol
Dmel\Smox
Species
D. melanogaster
Name
Smad on X
Annotation Symbol
CG2262
Feature Type
protein_coding_gene
FlyBase ID
FBgn0025800
Gene Model Status
Current
Stock Availability
18 publicly available
Gene Summary
Smad on X (Smox) encodes the primary transcription factor that mediates Activin signalling. Upon phosphorylation by the receptor encoded by babo, it forms a complex with the co-Smad protein encoded by Med. This complex translocates to the nucleus where it regulates expression of target genes. [Date last reviewed: 2018-10-18] (FlyBase Gene Snapshot)
All Summaries
Also Known As

dSmad2, Smad2, Smad, l(1)G0348, Sad

Key Links
Genomic Location
Cytogenetic map
7D14-7D15
Sequence location
Recombination map
1-22
RefSeq locus
NC_004354 REGION:8137041..8142279
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 (33 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
enables chromatin DNA binding
inferred from direct assay
(Hevia et al., 2017, Bai et al., 2013)
enables protein binding
inferred from physical interaction with FLYBASE:vis; FB:FBgn0033748
(Hyman et al., 2003)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
enables co-SMAD binding
traceable author statement
(Upadhyay et al., 2017)
enables DNA-binding transcription factor activity, RNA polymerase II-specific
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
enables I-SMAD binding
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
enables RNA polymerase II cis-regulatory region sequence-specific DNA binding
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
Biological Process (23 terms)
Terms Based on Experimental Evidence (19 terms)
CV Term
Evidence
References
involved_in activin receptor signaling pathway
inferred from mutant phenotype
(Langerak et al., 2018, Chng et al., 2014, Peterson et al., 2012)
inferred from genetic interaction with FLYBASE:babo; FB:FBgn0011300
(Lee et al., 2024, Wells et al., 2017)
inferred from direct assay
(Upadhyay et al., 2020, Gesualdi and Haerry, 2007, Parker et al., 2006, Lee-Hoeflich et al., 2005, Zheng et al., 2003, Brummel et al., 1999, Das et al., 1999)
inferred from genetic interaction with UniProtKB:A1Z7L8
(Zheng et al., 2003)
inferred from genetic interaction with FLYBASE:myo; FB:FBgn0026199
(Augustin et al., 2017)
inferred from genetic interaction with FLYBASE:Actβ; FB:FBgn0024913
(Hevia and de Celis, 2013)
involved_in axon guidance
inferred from mutant phenotype
(Parker et al., 2006)
involved_in cellular response to glucose stimulus
inferred from mutant phenotype
(Chng et al., 2014)
acts_upstream_of_positive_effect determination of adult lifespan
inferred from mutant phenotype
(Langerak et al., 2018)
involved_in glucose homeostasis
inferred from mutant phenotype
(Song et al., 2017)
involved_in imaginal disc growth
inferred from mutant phenotype
(Upadhyay et al., 2020)
involved_in imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Marquez et al., 2001)
involved_in mushroom body development
inferred from mutant phenotype
(Zheng et al., 2003)
involved_in negative regulation of autophagy
inferred from mutant phenotype
(Bai et al., 2013)
involved_in neuron development
inferred from mutant phenotype
(Zheng et al., 2006)
acts_upstream_of positive regulation of glycogen catabolic process
inferred from mutant phenotype
(Song et al., 2017)
involved_in positive regulation of imaginal disc growth
inferred from mutant phenotype
(Gesualdi and Haerry, 2007)
involved_in positive regulation of neuroblast proliferation
inferred from mutant phenotype
(Zhu et al., 2008)
involved_in positive regulation of transcription by RNA polymerase II
inferred from direct assay
(Das et al., 1999)
involved_in R8 cell fate specification
inferred from mutant phenotype
(Wells et al., 2017)
involved_in regulation of cell proliferation involved in imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Peterson and O'Connor, 2013)
involved_in regulation of imaginal disc-derived wing size
inferred from mutant phenotype
(Hevia and de Celis, 2013)
involved_in regulation of synapse structure or activity
inferred from mutant phenotype
(Kim and O'Connor, 2014)
involved_in SMAD protein signal transduction
inferred from direct assay
(Lee-Hoeflich et al., 2005)
Terms Based on Predictions or Assertions (7 terms)
CV Term
Evidence
References
involved_in activin receptor signaling pathway
inferred from biological aspect of ancestor with PANTHER:PTN001008908
(GO Reference Genome Project, 2011-)
involved_in anatomical structure morphogenesis
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
involved_in cell differentiation
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
involved_in dendrite morphogenesis
non-traceable author statement
(Grueber and Jan, 2004)
involved_in positive regulation of transcription by RNA polymerase II
inferred from biological aspect of ancestor with PANTHER:PTN001008908
(GO Reference Genome Project, 2011-)
involved_in regulation of DNA-templated transcription
(InterPro Project Members, 2004-)
involved_in SMAD protein signal transduction
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
Cellular Component (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
located_in cytosol
inferred from direct assay
(Hevia et al., 2017)
part_of heteromeric SMAD protein complex
inferred from physical interaction with FLYBASE:Med; FB:FBgn0288966
(Brummel et al., 1999, Das et al., 1999)
located_in nucleus
inferred from direct assay
(Hevia et al., 2017)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
part_of heteromeric SMAD protein complex
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
part_of transcription regulator complex
inferred from electronic annotation with InterPro:IPR013019
(InterPro Project Members, 2004-)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
Belongs to the dwarfin/SMAD family. (O96660)
Protein Signatures (InterPro)
Summaries
Gene Snapshot
Smad on X (Smox) encodes the primary transcription factor that mediates Activin signalling. Upon phosphorylation by the receptor encoded by babo, it forms a complex with the co-Smad protein encoded by Med. This complex translocates to the nucleus where it regulates expression of target genes. [Date last reviewed: 2018-10-18]
Gene Group (FlyBase)
MAD HOMOLOGY DOMAIN TRANSCRIPTION FACTORS -
The Mother against Dpp (MAD) homology (MH) domain transcription factors are sequence-specific DNA-binding proteins that regulate transcription. These members contain an N-terminal sequence-specific DNA-binding MH1 and a C-terminal MH2 domain that mediates the formation of oligomeric complexes. (Adapted from FBrf0208242 and FBrf0206210).
Pathway (FlyBase)
ACTIVIN SIGNALING PATHWAY CORE COMPONENTS -
The activin signaling pathway is one of two branches of Transforming Growth Factor-β family signaling in Drosophila. The binding of an activin family dimer to a heterodimeric serine/threonine kinase receptor complex results in the phosphorylation of Smox, a members of the Smad family. Smox forms a complex with the co-Smad, Med. This complex translocates into the nucleus and regulates the transcription of target genes in concert with other nuclear cofactors. (Adapted from FBrf0236482.)
Protein Function (UniProtKB)
Receptor-regulated SMAD (R-SMAD) involved in the activin-like TGF-beta pathway (PubMed:10320478). Acts downstream of the type I transmembrane receptor serine/threonine kinase babo (PubMed:10320478). Competitively regulates phosphorylation of Mad, the R-SMAD of the BMP signaling pathway, by babo (PubMed:22563507). Heterodimerizes with the Co-SMAD Med/medea upon activation of the activin pathway and translocates to the nucleus, where it transcriptionally activates activin-responsive genes (PubMed:10320478). Involved in axonal and dendritic pruning and neural remodeling of mushroom body gamma neurons during metamorphosis, probably through regulating expression of the ecdysteroid receptor EcR (PubMed:12581521). Involved in differentiation of dorsal cluster and ellipsoid body adult-specific neurons prior to metamorphosis (PubMed:16437159). Required for axon guidance or outgrowth of ISNb and SNa axons (PubMed:17119022). Involved in regulation of wing imaginal disc growth (PubMed:18820452). Signaling by daw or myo is involved in lifespan determination through regulation of proteasome-mediated degradation of misfolded or damaged proteins (PubMed:29615416).
(UniProt, O96660)
Summary (Interactive Fly)

Dpp signaling pathway - Smad2 and Smad3 homolog - a target of Baboon - Activin signalling is required for ribosome biogenesis and cell growth in Drosophila salivary glands

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

Please see the JBrowse view of Dmel\Smox 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 O96660)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.50

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Smox-RA
FBtr0071192
NM_078524
3171
486
Smox-RB
FBtr0340159
NM_001298077
4521
486
Additional Transcript Data and Comments
Reported size (kB)

~4.5 (northern blot)

(Henderson and Andrew, 1998)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
Smox-PA
FBpp0071140
53.7
486
7.63
O96660
NP_511079
AAF46330
Smox-PB
FBpp0309145
53.7
486
7.63
O96660
NP_001285006
AHN59477
Polypeptides with Identical Sequences

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

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

486 (aa)

(Brummel et al., 1999)

484 (aa)

(Henderson and Andrew, 1998)
Comments

Activated babo protein phosphorylates the C-terminal SSXS motif of Smox protein. Med and Smox proteins form some complexes in the absence of activated babo protein, but the level of complex formation increases significantly in the presence of the activated babo protein. Smox protein also interacts transiently with putFBtr0091350:pb-XRbabo protein complexes.

(Brummel et al., 1999)
External Data
Subunit Structure (UniProtKB)

Associates with a receptor serine/threonine kinase complex composed of put and babo; dissociates upon being phosphorylated by babo (PubMed:10320478, PubMed:9887103). Interacts (when phosphorylated on the SSXS motif) with Med/medea; the interaction is dependent on a receptor serine/threonine kinase complex (PubMed:10320478, PubMed:16951053, PubMed:9887103). Heterodimerizes with the Co-SMAD Med/medea upon activation of the activin pathway and translocates to the nucleus, where it transcriptionally activates activin-responsive genes (PubMed:10320478).

(UniProt, O96660)
Post Translational Modification

Phosphorylated in response to Actbeta/activin and daw signaling (PubMed:12581521, PubMed:18820452, PubMed:19766717). Phosphorylated on a C-terminal SSXS motif by the receptor serine/threonine kinase complex composed of the type I receptor babo and type II receptor put/punt (PubMed:10320478, PubMed:18820452, PubMed:19766717, PubMed:22563507, PubMed:9887103). Phosphorylation by babo induces heterodimerization with med/medea and nuclear localization (PubMed:10320478). Phosphorylation by babo increases protein turnover (PubMed:22563507).

(UniProt, O96660)
Crossreferences
InterPro - A database of protein families, domains and functional sites
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\Smox 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).

-1.45

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Henderson and Andrew, 1998)
embryonic stage -- adult stage
(Henderson and Andrew, 1998)
pre-blastoderm stage
organism
(Brummel et al., 1999)
embryonic stage 1 -- 2
pole plasm
(Yatsu et al., 2008)
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
organism | ubiquitous
(Fisher et al., 2012)
primordial germ cell
(Fisher et al., 2012)
embryonic stage 6 -- 17
mesoderm
(Brummel et al., 1999)
embryonic stage 7 -- 8
organism | ubiquitous
(Fisher et al., 2012)
primordial germ cell
(Fisher et al., 2012)
embryonic stage 9 -- 10
germline cell
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 11 -- 12
germline cell
(Fisher et al., 2012)
organism | faint | ubiquitous
(Fisher et al., 2012)
embryonic stage 13 -- 16
dorsal pharyngeal muscle cell
(Fisher et al., 2012)
embryonic anal pad
(Fisher et al., 2012)
embryonic brain
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
larval somatic muscle cell
(Fisher et al., 2012)
larval visceral muscle cell
(Fisher et al., 2012)
gonad
(Fisher et al., 2012)
organism | faint | ubiquitous
(Fisher et al., 2012)
larval ventral nerve cord
(Fisher et al., 2012)
third instar larval stage
larval brain | restricted
(Brummel et al., 1999)
imaginal disc
(Brummel et al., 1999)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage -- adult stage
(Henderson and Andrew, 1998)
embryonic stage 1 -- 4

Comment: reference states 0-2 hr AEL

(Henderson and Andrew, 1998)
adult stage | female
(Henderson and Andrew, 1998)
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytosol
inferred from direct assay
(Hevia et al., 2017)
part_of heteromeric SMAD protein complex
inferred from physical interaction with FLYBASE:Med; FB:FBgn0288966
(Brummel et al., 1999, Das et al., 1999)
located_in nucleus
inferred from direct assay
(Hevia et al., 2017)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Smox 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
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 13 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 24 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Smox
Transgenic constructs containing regulatory region of Smox
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 | larval stage | recessive
lethal | P-stage | recessive
lethal - all die before end of P-stage
lethal - all die before end of P-stage, with Scer\GAL4da.G32
lethal - all die before end of pupal stage, with Scer\GAL4pnr-MD237
lethal - all die during P-stage, with Scer\GAL4Mef2.PR
partially lethal - majority die | recessive
viable (with Dp(1;3)DC185)
viable (with Dp(1;3)DC186)
Sterility
Allele
fertile (with Dp(1;3)DC185)
fertile (with Dp(1;3)DC186)
Other Phenotypes
Allele
abnormal cell growth, with Scer\GAL4A9
abnormal flight | progressive, with Scer\GAL4Mhc.PW
abnormal neuroanatomy
abnormal neuroanatomy | embryonic stage
abnormal neuroanatomy | embryonic stage | germline clone
abnormal neuroanatomy | larval stage
abnormal neuroanatomy | somatic clone
abnormal size | adult stage, with Scer\GAL4AB1
abnormal size | adult stage, with Scer\GAL4Dll-md23
abnormal size | adult stage, with Scer\GAL4nub.PU
abnormal size | adult stage | somatic clone
abnormal size | larval stage
abnormal size | late third instar larval stage
abnormal size | third instar larval stage
abnormal size | third instar larval stage, with Scer\GAL432B
abnormal size | third instar larval stage, with Scer\GAL4da.G32
abnormal size | third instar larval stage, with Scer\GAL4esg-05730B
abnormal size | third instar larval stage, with Scer\GAL4nub-AC-62, Scer\GAL4tsh-md621
abnormal size | third instar larval stage, with Scer\GAL4sd-SG29.1
decreased cell number | adult stage, with Scer\GAL4nub.PU
decreased cell number | progressive | spermatogenesis, with Scer\GAL4upd1-E132
decreased cell number | third instar larval stage
decreased cell size | adult stage, with Scer\GAL4nub.PU
decreased cell size | late third instar larval stage
decreased occurrence of cell division | third instar larval stage
decreased size | larval stage
decreased size | pharate adult stage, with Scer\GAL4da.G32
increased cell number | adult stage, with Scer\GAL4nub.PU
increased cell number | late third instar larval stage
increased cell size | adult stage, with Scer\GAL4nub.PU
increased size | larval stage
increased size | larval stage, with Scer\GAL4NP5130
long lived, with Scer\GAL4Mhc.PW
long lived | RU486 conditional, with Scer\GAL4αTub84B.Switch.PK
short lived | male
some die during pharate adult stage
some die during pupal stage, with Scer\GAL4pnr-MD237
visible, with Scer\GAL4A9
visible, with Scer\GAL4vg.MQ.Exel
Phenotype manifest in
Allele
adult brain | pharate adult stage, with Scer\GAL4da.G32
adult medulla neuropil associated glial cell
adult optic lobe | pharate adult stage, with Scer\GAL4da.G32
adult optic lobe | somatic clone
adult somatic muscle cell | progressive, with Scer\GAL4Mhc.PW
autophagosome, with Scer\GAL4Mhc.PW
axon & eye photoreceptor cell
chaeta
dorsal cluster neuron & axon | somatic clone
dorsal cluster neuron & neurite | somatic clone
embryonic/larval anal pad | larval stage
eye photoreceptor cell | larval stage
fascicle | somatic clone
gamma-lobe & dendrite | somatic clone
gamma-lobe & neuron | somatic clone
hub cell | decreased number, with Scer\GAL4upd1-E132
lamina
lamina anlage glial cell
larval abdominal segmental nerve | germline clone
larval brain
larval brain | late third instar larval stage
larval inner optic anlage
larval outer optic anlage
leg, with Scer\GAL4Dll-md23
leg | somatic clone
lobula columnar neuron LC14 | somatic clone
lysosome | progressive, with Scer\GAL4Mhc.PW
medulla anlage
nucleus | adult stage, with Scer\GAL4AB1
photoreceptor cell R7 & axon
photoreceptor cell R7 | somatic clone
photoreceptor cell R8 & axon
protocerebrum | somatic clone
salivary gland | adult stage, with Scer\GAL4AB1
transmedullary neuron Tm20 | somatic clone
wing
wing, with Scer\GAL4A9
wing, with Scer\GAL4nub.PU
wing, with Scer\GAL4nub-AC-62
wing | somatic clone
wing blade, with Scer\GAL4vg.MQ.Exel
wing disc | larval stage
wing disc | larval stage, with Scer\GAL4da.G32
wing disc | larval stage | somatic clone
wing disc | late third instar larval stage
wing disc | third instar larval stage, with Scer\GAL432B
wing disc | third instar larval stage, with Scer\GAL4esg-05730B
wing disc | third instar larval stage, with Scer\GAL4nub-AC-62, Scer\GAL4tsh-md621
wing disc | third instar larval stage, with Scer\GAL4sd-SG29.1
wing pouch | larval stage, with Scer\GAL4NP5130
wing vein | ectopic, with Scer\GAL4nub.PU
wing vein | ectopic | 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) (8)
Hsap\SMAD3
11 of 14
Yes
Yes
1  
Hsap\SMAD2
10 of 14
No
Yes
1  
Hsap\SMAD1
6 of 14
No
No
1  
Hsap\SMAD5
6 of 14
No
No
2  
Hsap\SMAD9
6 of 14
No
No
Hsap\SMAD4
2 of 14
No
No
8  
Hsap\SMAD6
2 of 14
No
No
2  
Hsap\SMAD7
2 of 14
No
No
1  
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (8)
Rnor\Smad3
11 of 14
Yes
Yes
Rnor\Smad2
10 of 14
No
Yes
Rnor\Smad1
6 of 14
No
No
Rnor\Smad5
6 of 14
No
No
Rnor\Smad9
5 of 14
No
No
Rnor\Smad6
2 of 14
No
No
Rnor\Smad7
2 of 14
No
No
Rnor\Smad4
1 of 14
No
No
Mus musculus (laboratory mouse) (8)
Mmus\Smad3
11 of 14
Yes
Yes
Mmus\Smad2
10 of 14
No
Yes
Mmus\Smad1
6 of 14
No
No
Mmus\Smad5
6 of 14
No
No
Mmus\Smad9
5 of 14
No
No
Mmus\Smad4
2 of 14
No
No
Mmus\Smad6
2 of 14
No
No
Mmus\Smad7
2 of 14
No
No
Xenopus tropicalis (Western clawed frog) (9)
Xtro\smad2
8 of 13
Yes
Yes
Xtro\smad3
7 of 13
No
Yes
Xtro\cdk6
5 of 13
No
No
Xtro\smad1
5 of 13
No
No
Xtro\smad9
5 of 13
No
No
Xtro\tmem54
4 of 13
No
No
Xtro\rpl14
1 of 13
No
No
Xtro\smad7
1 of 13
No
No
Xtro\tspan15
1 of 13
No
No
Danio rerio (Zebrafish) (13)
Drer\smad3b
12 of 14
Yes
Yes
Drer\smad2
11 of 14
No
Yes
Drer\smad3a
11 of 14
No
Yes
Drer\smad5
6 of 14
No
No
Drer\smad1
5 of 14
No
No
Drer\smad9
5 of 14
No
No
Drer\smad4a
2 of 14
No
No
Drer\smad6a
2 of 14
No
No
Drer\smad6b
2 of 14
No
No
Drer\smad7
2 of 14
No
No
Drer\smad10a
2 of 14
No
No
Drer\gb:cr929477
1 of 14
No
No
Drer\smad4b
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (8)
Cele\sma-2
8 of 14
Yes
No
Cele\daf-8
4 of 14
No
Yes
Cele\sma-3
4 of 14
No
Yes
Cele\daf-14
2 of 14
No
Yes
Cele\tag-68
2 of 14
No
No
Cele\C34E11.2
1 of 14
No
No
Cele\daf-3
1 of 14
No
No
Cele\sma-4
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (4)
Agam\AgaP_AGAP009777
12 of 12
Yes
Yes
Agam\AgaP_AGAP008551
5 of 12
No
No
Agam\AgaP_AGAP002902
3 of 12
No
No
Agam\AgaP_AGAP007416
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:Smox. 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) (5)
Mad
9 of 13
Dad
4 of 13
Med
4 of 13
exp
1 of 13
reb
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 2 )
    Modifiers Based on Experimental Evidence ( 4 )
    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?
    SMAD3; SMAD family member 3
    11 of 14
    603109
     
    SMAD2; SMAD family member 2
    10 of 14
    601366
     
    SMAD1; SMAD family member 1
    6 of 14
    601595
         
        SMAD5; SMAD family member 5
        6 of 14
        603110
             
            SMAD9; SMAD family member 9
            6 of 14
            603295
            Functional Complementation Data
            Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
            Interactions
            Summary of Physical Interactions
            Summary of Genetic Interactions
            Interaction Browsers
            View in FlyBase Interactions Browser   View in MIST tool (external link)
            Please look at the allele data for full details of the genetic interactions
            Starting gene(s)
            Interaction type
            Interacting gene(s)
            Reference
            Smox
            suppressible
            Acf
            (Hevia et al., 2017)
            Smox
            enhanceable
            Actβ
            (Gesualdi and Haerry, 2007)
            Smox
            suppressible
            babo
            (Upadhyay et al., 2020, Peterson and O'Connor, 2013)
            Smox
            enhanceable
            babo
            (Gesualdi and Haerry, 2007)
            Smox
            suppressible
            brat
            (Hevia et al., 2017)
            Smox
            suppressible
            bun
            (Hevia et al., 2017)
            Smox
            suppressible
            CG3649
            (Hevia et al., 2017)
            Smox
            suppressible
            CG8420
            (Hevia et al., 2017)
            Smox
            suppressible
            CG9008
            (Hevia et al., 2017)
            Smox
            suppressible
            CG9733
            (Hevia et al., 2017)
            Smox
            suppressible
            CG10089
            (Hevia et al., 2017)
            Smox
            suppressible
            CG12182
            (Hevia et al., 2017)
            Smox
            suppressible
            CG13083
            (Hevia et al., 2017)
            Smox
            suppressible
            CG14692
            (Hevia et al., 2017)
            Smox
            suppressible
            CG15152
            (Hevia et al., 2017)
            Smox
            suppressible
            CG15212
            (Hevia et al., 2017)
            Smox
            suppressible
            CG18473
            (Hevia et al., 2017)
            Smox
            suppressible
            CG31436
            (Hevia et al., 2017)
            Smox
            suppressible
            Cluap1
            (Hevia et al., 2017)
            Smox
            suppressible
            Cpr57A
            (Hevia et al., 2017)
            Smox
            suppressible
            Cpr67Fa2
            (Hevia et al., 2017)
            Smox
            suppressible
            CR6434
            (Hevia et al., 2017)
            Smox
            enhanceable
            CycE
            (Hevia et al., 2017)
            Smox
            suppressible
            CycE
            (Hevia et al., 2017)
            Smox
            suppressible
            Cyp9c1
            (Hevia et al., 2017)
            Smox
            suppressible
            Cyp310a1
            (Hevia et al., 2017)
            Smox
            suppressible
            daw
            (Gesualdi and Haerry, 2007)
            Smox
            suppressible
            DJ-1α
            (Hevia et al., 2017)
            Smox
            suppressible
            GstE8
            (Hevia et al., 2017)
            Smox
            suppressible
            hpo
            (Hevia et al., 2017)
            Smox
            suppressible
            InR
            (Hevia et al., 2017)
            Smox
            enhanceable
            InR
            (Hevia et al., 2017)
            Smox
            enhanceable
            lectin-28C
            (Hevia et al., 2017)
            Smox
            suppressible
            Mad
            (Hevia et al., 2017)
            Smox
            suppressible
            meng
            (Hevia et al., 2017)
            Smox
            suppressible
            mod(mdg4)
            (Hevia et al., 2017)
            Smox
            suppressible
            myo
            (Upadhyay et al., 2020, Gesualdi and Haerry, 2007)
            Smox
            suppressible
            NetA
            (Hevia et al., 2017)
            Smox
            suppressible
            Ofut1
            (Hevia et al., 2017)
            Smox
            suppressible
            Optix
            (Hevia et al., 2017)
            Smox
            suppressible
            Oseg5
            (Hevia et al., 2017)
            Smox
            suppressible
            Pih1D1
            (Hevia et al., 2017)
            Smox
            suppressible
            Pten
            (Hevia et al., 2017)
            Smox
            suppressible
            Rala
            (Hevia et al., 2017)
            Smox
            enhanceable
            RfC3
            (Hevia et al., 2017)
            Smox
            suppressible
            rpk
            (Hevia et al., 2017)
            Smox
            suppressible
            Spn100A
            (Hevia et al., 2017)
            Smox
            suppressible
            Srr
            (Hevia et al., 2017)
            Smox
            enhanceable
            stg
            (Hevia et al., 2017)
            Smox
            suppressible
            stg
            (Hevia et al., 2017)
            Smox
            suppressible
            Thg
            (Hevia et al., 2017)
            Smox
            suppressible
            Vajk2
            (Hevia et al., 2017)
            Smox
            suppressible
            Vha16-5
            (Hevia et al., 2017)
            Smox
            enhanceable
            wb
            (Hevia et al., 2017)
            Smox
            suppressible
            yki
            (Hevia et al., 2017)
            Smox
            enhanceable
            yki
            (Hevia et al., 2017)
            Starting gene(s)
            Interaction type
            Interacting gene(s)
            Reference
            babo
            enhanceable
            Smox
            (Peterson et al., 2012, Gesualdi and Haerry, 2007)
            babo
            suppressible
            Smox
            (Peterson et al., 2012)
            hpo
            suppressible
            Smox
            (Hevia et al., 2017)
            InR
            enhanceable
            Smox
            (Hevia et al., 2017)
            InR
            suppressible
            Smox
            (Hevia et al., 2017)
            myo
            suppressible
            Smox
            (Augustin et al., 2017)
            plum
            suppressible
            Smox
            (Yu et al., 2013)
            Pten
            suppressible
            Smox
            (Hevia et al., 2017)
            Scgδ
            suppressible
            Smox
            (Goldstein et al., 2011)
            Snr1
            suppressible
            Smox
            (Tegeder et al., 2019)
            yki
            enhanceable
            Smox
            (Hevia et al., 2017)
            External Data
            Subunit Structure (UniProtKB)
            Associates with a receptor serine/threonine kinase complex composed of put and babo; dissociates upon being phosphorylated by babo (PubMed:10320478, PubMed:9887103). Interacts (when phosphorylated on the SSXS motif) with Med/medea; the interaction is dependent on a receptor serine/threonine kinase complex (PubMed:10320478, PubMed:16951053, PubMed:9887103). Heterodimerizes with the Co-SMAD Med/medea upon activation of the activin pathway and translocates to the nucleus, where it transcriptionally activates activin-responsive genes (PubMed:10320478).
            (UniProt, O96660 )
            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
            Class of Gene
            Genomic Location and Detailed Mapping Data
            Chromosome (arm)
            X
            Recombination map
            1-22
            Cytogenetic map
            7D14-7D15
            Sequence location
            FlyBase Computed Cytological Location
            Cytogenetic map
            Evidence for location
            7D14-7D15
            Limits computationally determined from genome sequence between P{EP}Ubc-E2HEP1303 and P{EP}CG1583EP1516&P{EP}CG1583EP325
            Experimentally Determined Cytological Location
            Cytogenetic map
            Notes
            References
            7D-7D
            (determined by in situ hybridisation)
            (Peter et al., 2002, Schäfer et al., 1999.12.2, Henderson and Andrew, 1998)
            7D10-7D12
            (Zheng et al., 2003)
            7D7-7D16
            (Andrew, 1998.3.10, Henderson, 1998.7.15)
            7D-7D
            (Brummel, 1998.10.26, Das, 1998.11.23)
            7D6-7D17
            (Das et al., 1999)
            Mapped to 7D6--7D17 using grids of P1 clones.
            (Das et al., 1999)
            Experimentally Determined Recombination Data
            Location

            1-22

            (FlyBase, 2016)

            1-22.3

            (Comeron et al., 2012)
            Left of (cM)
            Right of (cM)
            Notes
            Stocks and Reagents
            Stocks (18)
            Genomic Clones (11)
             

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

            cDNA Clones (62)
             

            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

            (Bai et al., 2013)
            Commercially Available Antibodies
             
            Cell Line Information
            Publicly Available Cell Lines
             
              Other Stable Cell Lines
               
                Other Comments

                Smox is required during the late larval stage for development of adult-specific neurons.

                (Zheng et al., 2006)

                babo/Smox mediated TGFΒ/Activin signalling mediates remodeling of mushroom body neurons via controlling EcR-B1 expression.

                (Zheng et al., 2003)

                Overexpression of Med, Smox, Hsap\MADH2 or Hsap\MADH4 causes a similar phenotype in the wing.

                (Marquez et al., 2001)

                Smox is highly expressed in tissues that require babo function and can be phosphorylated by overexpression of activated babo or wild type babo and put together.

                (Brummel et al., 1999)
                Relationship to Other Genes
                Source for database merge of

                Source for merge of: Smox Sad

                (Gelbart, 2000.4.17)

                Source for merge of: Smox tmp

                (Padgett, 2000.8.24)
                Additional comments

                The Smox gene may have been derived from the Mad gene by retroposition.

                (Langille and Clark, 2007)
                Nomenclature History
                Source for database identify of

                Source for identity of: Smox CG2262

                (Zhang, 1999.11)
                Nomenclature comments
                Etymology
                Synonyms and Secondary IDs (26)
                Reported As
                Symbol Synonym
                CG2262
                (Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Perkins et al., 2009.3.31)
                DSMAD2
                (Tanimoto et al., 2000)
                DSmad2
                (Pistillo and Desplan, 2007, Pistillo et al., 2007)
                SMAD2
                (Hunt and Mannervik, 2024, Mattila and Hietakangas, 2017, Sanyal et al., 2004)
                SMOX
                (Fan et al., 2018, Liu and Jin, 2017, Ayyaz et al., 2015, Taylor-Harding et al., 2004)
                Sad
                (Andrew, 1998.3.10)
                Smad
                (Zhang and Cai, 2020, Hevia and de Celis, 2013, Peterson and O'Connor, 2013)
                Smad2
                (Kim and O'Connor, 2021, Sood et al., 2021, Texada et al., 2020, Setiawan et al., 2018, Augustin et al., 2017, Chng et al., 2017, Hevia et al., 2017, Upadhyay et al., 2017, Di Cara and King-Jones, 2016, Niwa and Niwa, 2016, Ting et al., 2014, Peterson and O'Connor, 2013, Yamanaka et al., 2013, Arora et al., 2008, Kamiya et al., 2008, Zhu et al., 2008, Parker et al., 2006, Serpe and O'Connor, 2006, Yang et al., 2004, Padgett, 1999)
                Smad2/3
                (van Soldt and Cardoso, 2020)
                SmoX
                (Yu et al., 2013, Yu et al., 2013)
                Smox
                (Lauraine et al., 2024, Bastin and Eleftherianos, 2023, Lassetter et al., 2023, Saha et al., 2022, Bilder et al., 2021, Fabian et al., 2021, Huang et al., 2021, Kramer et al., 2021, Cui et al., 2020, Graves et al., 2020, Han et al., 2020, Jasper, 2020, Li et al., 2020, Zhang et al., 2020, Shokri et al., 2019, Tegeder et al., 2019, Bischof et al., 2018, Langerak et al., 2018, Muñoz-Soriano et al., 2018, Song et al., 2018, Stephano et al., 2018, Transgenic RNAi Project members, 2017-, Zhu et al., 2017, Bonfini et al., 2016, Hu et al., 2016, Ayyaz et al., 2015, Lengil et al., 2015, Mattila et al., 2015, Schertel et al., 2015, dos Santos, 2014, Hsiao et al., 2014, Bai et al., 2013, Chintapalli et al., 2013, Marinho et al., 2013, Spokony and White, 2013.5.10, Yamamoto et al., 2013-, Hazelett et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Sorrentino et al., 2012, Zoller and Schulz, 2012, Goldstein et al., 2011, Toku et al., 2011, Blanco et al., 2010, Nedelsky et al., 2010, Popodi et al., 2010-, Venken et al., 2010, Jensen et al., 2009, Batut et al., 2008, Kamiya et al., 2008, Yatsu et al., 2008, Beltran et al., 2007, Langille and Clark, 2007, Li et al., 2007, Dudu, 2006, Kulkarni et al., 2006)
                dSMAD2
                (Bickel et al., 2008, Gesualdi and Haerry, 2007, Pinto et al., 2006, Massagué and Chen, 2000)
                dSmad
                (Furusawa and Emoto, 2021)
                dSmad2
                (Castro et al., 2024, Kimura et al., 2024, Lee et al., 2024, Shin et al., 2024, Truman and Riddiford, 2023, Pan et al., 2020, Upadhyay et al., 2020, Moss-Taylor et al., 2019, Wang et al., 2019, Zhu et al., 2019, Anderson and Wharton, 2017, Makhijani et al., 2017, Song et al., 2017, Wells et al., 2017, Van Bortle et al., 2015, Ting et al., 2014, Kniss et al., 2013, Peterson and O'Connor, 2013, Peterson et al., 2012, Smith et al., 2012, Sorrentino et al., 2012, Takaesu et al., 2012, Hadjieconomou et al., 2011, Quijano et al., 2011, Ellis et al., 2010, Quijano et al., 2010, Sander et al., 2010, Batut et al., 2008, Kamiya et al., 2008, Ng, 2008, Pinto et al., 2008, Barrio et al., 2007, Newfeld et al., 2007, Ting et al., 2007, Takaesu et al., 2006, Zheng et al., 2006, Zhu et al., 2006, Lee-Hoeflich et al., 2005, Marques, 2005, Massagué et al., 2005, Funakoshi et al., 2004, Grueber and Jan, 2004, Koh et al., 2004, Hyman et al., 2003, Liang et al., 2003, Zheng et al., 2003, Zheng et al., 2003, Padgett et al., 2002, Parker and Arora, 2002, Marquez et al., 2001, Padgett et al., 2001, Stefancsik et al., 2001, Stefancsik et al., 2001, Attisano and Wrana, 2000, Das et al., 2000, Massague and Wotton, 2000, Nakayama et al., 2000, Newfeld et al., 2000, Zimmerman and Padgett, 2000, Brummel et al., 1999, Das et al., 1999, Das et al., 1999, Raftery and Sutherland, 1999, Zhang and Derynck, 1999)
                dsmad2
                (Ayyaz et al., 2015, Ting et al., 2014, Ting et al., 2014, Ting et al., 2007)
                l(1)G0348
                (Schäfer et al., 1999.12.2)
                sad
                (Henderson and Andrew, 1998)
                smad2
                (Ell et al., 2024, Brummel, 1998.10.26)
                smox
                (Deng et al., 2022, Herrera et al., 2021, Cai et al., 2019, Whittle and Extavour, 2019, Jeibmann et al., 2017, Baëza et al., 2015, Peterson et al., 2012, Stronach and Perrimon, 2002)
                ted
                (Padgett et al., 1998)
                tmp
                Name Synonyms
                Sisters against dpp
                Smad on X
                (Graves et al., 2020, Marinho et al., 2013, Zoller and Schulz, 2012, Henderson and Andrew, 1998)
                Smox/dSmad2
                (Li et al., 2007)
                sisters against dpp
                (Henderson and Andrew, 1998)
                temporarily deranged
                (Padgett et al., 1998)
                Secondary FlyBase IDs
                • FBgn0024186
                • FBgn0024304
                • FBgn0040171
                Datasets (0)
                Study focus (0)
                Experimental Role
                Project
                Project Type
                Title
                Study result (0)
                Result
                Result Type
                Title
                External Crossreferences and Linkouts ( 68 )
                Sequence Crossreferences
                NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
                GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
                GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
                RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
                UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
                UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
                UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
                Other crossreferences
                AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
                BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
                DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
                EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
                FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
                FlyMine - An integrated database for Drosophila genomics
                InterPro - A database of protein families, domains and functional sites
                KEGG Genes - Molecular building blocks of life in the genomic space.
                MARRVEL_MODEL - MARRVEL (model organism gene)
                Linkouts
                BioGRID - A database of protein and genetic interactions.
                Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
                DroID - A comprehensive database of gene and protein interactions.
                DRSC - Results frm RNAi screens
                Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
                FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
                FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
                Flygut - An atlas of the Drosophila adult midgut
                FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
                iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
                Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                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 (239)