FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\Med
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General Information
Symbol
Dmel\Med
Species
D. melanogaster
Name
Medea
Annotation Symbol
CG1775
Feature Type
protein_coding_gene
FlyBase ID
FBgn0288966
Gene Model Status
Current
Stock Availability
23 publicly available
Gene Summary
Medea (Med) encodes a protein that belongs to the highly conserved Smad family. It can bind its siblings encoded by Mad or Smox to facilitate signal transduction for the product of dpp or Activin ligands in the TGF-beta family. Med-complexes function as transcriptional regulators. Many developmental roles include dorsal-ventral patterning, patterning and proliferation of the wing disc and gene expression in the mushroom body of the larval brain. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
All Summaries
Also Known As

l(3)SG70, dSmad4, Smad4, l(3)11m-254, demented

Key Links
Genomic Location
Cytogenetic map
100C7-100D1
Sequence location
Recombination map
3-102
RefSeq locus
NT_033777 REGION:31611046..31615375
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 (36 terms)
Molecular Function (7 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
enables DNA-binding transcription activator activity, RNA polymerase II-specific
inferred from direct assay
(Vuilleumier et al., 2022, Weiss et al., 2010)
enables DNA-binding transcription repressor activity, RNA polymerase II-specific
inferred from direct assay
(Yao et al., 2006, Muller et al., 2003)
enables protein binding
inferred from physical interaction with UniProtKB:Q9NB71
(McCabe et al., 2004)
enables RNA polymerase II cis-regulatory region sequence-specific DNA binding
inferred from direct assay
(Xu et al., 1998)
enables RNA polymerase II transcription regulatory region sequence-specific DNA binding
inferred from direct assay
(Weiss et al., 2010)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
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 (17 terms)
CV Term
Evidence
References
involved_in activin receptor signaling pathway
inferred from direct assay
(Brummel et al., 1999, Das et al., 1999)
involved_in BMP signaling pathway
inferred from direct assay
(Vuilleumier et al., 2022, Weiss et al., 2010, Kamiya et al., 2008, Yao et al., 2006, Muller et al., 2003, Dai et al., 2000, Das et al., 1998, Inoue et al., 1998)
inferred from genetic interaction with FLYBASE:dpp; FB:FBgn0000490
(Hudson et al., 1998, Wisotzkey et al., 1998, Raftery et al., 1995)
inferred from mutant phenotype
(Zhang et al., 2025, Stinchfield et al., 2012, McCabe et al., 2004, Hudson et al., 1998)
inferred from genetic interaction with FLYBASE:dpp; FB:FBgn0000490,FLYBASE:sax; FB:FBgn0003317
(Das et al., 1998)
inferred from mutant phenotype
(Raftery and Wisotzkey, 1996)
inferred from genetic interaction with FLYBASE:tkv; FB:FBgn0003716
(Hudson et al., 1998)
inferred from genetic interaction with FLYBASE:zen; FB:FBgn0004053
(Hudson et al., 1998)
involved_in compound eye morphogenesis
inferred from mutant phenotype
(Das et al., 1998)
involved_in dorsal/ventral axis specification
inferred from mutant phenotype
(Hudson et al., 1998)
involved_in dorsal/ventral pattern formation
inferred from mutant phenotype
(Das et al., 1998, Wisotzkey et al., 1998)
involved_in follicle cell of egg chamber development
inferred from mutant phenotype
(Kirilly et al., 2005)
involved_in germ-line stem cell division
inferred from mutant phenotype
(Xie and Spradling, 1998)
involved_in germ-line stem cell population maintenance
inferred from mutant phenotype
(Xie and Spradling, 1998)
involved_in imaginal disc-derived leg morphogenesis
inferred from mutant phenotype
(Marquez et al., 2001)
involved_in imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Marquez et al., 2001)
involved_in negative regulation of transcription by RNA polymerase II
inferred from direct assay
(Yao et al., 2006, Muller et al., 2003)
involved_in neuron development
inferred from mutant phenotype
(Zheng et al., 2006)
involved_in positive regulation of synaptic assembly at neuromuscular junction
inferred from mutant phenotype
(McCabe et al., 2004)
involved_in positive regulation of transcription by RNA polymerase II
inferred from direct assay
(Weiss et al., 2010, Dai et al., 2000, Das et al., 1999)
inferred from mutant phenotype
(Xu et al., 1998)
involved_in retinal ganglion cell axon guidance
inferred from mutant phenotype
(Yoshida et al., 2005)
involved_in SMAD protein signal transduction
inferred from direct assay
(Das et al., 1998)
involved_in somatic stem cell population maintenance
inferred from mutant phenotype
(Kirilly et al., 2005)
Terms Based on Predictions or Assertions (8 terms)
CV Term
Evidence
References
involved_in anatomical structure morphogenesis
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
involved_in BMP signaling pathway
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 heart development
traceable author statement
(Cripps and Olson, 2002, Chen and Fishman, 2000)
involved_in negative regulation of salivary gland boundary specification
traceable author statement
(Bradley et al., 2001)
involved_in regulation of DNA-templated transcription
(InterPro Project Members, 2004-)
involved_in regulation of transcription by RNA polymerase II
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
involved_in SMAD protein signal transduction
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
Cellular Component (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
located_in cytoplasm
inferred from high throughput direct assay
(Sarov et al., 2016)
located_in cytosol
inferred from direct assay
(Das et al., 1998, Wisotzkey et al., 1998)
part_of heteromeric SMAD protein complex
inferred from physical interaction with FLYBASE:Smox; FB:FBgn0025800
(Brummel et al., 1999, Das et al., 1999)
inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648
(Kamiya et al., 2008, Wisotzkey et al., 1998)
inferred from direct assay
(Inoue et al., 1998)
located_in nucleus
inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648
(Wisotzkey et al., 1998)
inferred from direct assay
(Das et al., 1998)
part_of RNA polymerase II transcription repressor complex
inferred from physical interaction with FLYBASE:shn; FB:FBgn0003396,FLYBASE:Mad; FB:FBgn0011648
(Yao et al., 2006, Muller et al., 2003)
part_of transcription regulator complex
inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648
(Weiss et al., 2010)
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. (O62609)
Protein Signatures (InterPro)
Summaries
Gene Snapshot
Medea (Med) encodes a protein that belongs to the highly conserved Smad family. It can bind its siblings encoded by Mad or Smox to facilitate signal transduction for the product of dpp or Activin ligands in the TGF-beta family. Med-complexes function as transcriptional regulators. Many developmental roles include dorsal-ventral patterning, patterning and proliferation of the wing disc and gene expression in the mushroom body of the larval brain. [Date last reviewed: 2019-03-14]
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)
BMP SIGNALING PATHWAY CORE COMPONENTS -
The Bone Morphogenetic Protein (BMP) signaling pathway is one of two branches of Transforming Growth Factor-β family signaling in Drosophila. The binding of a BMP family dimer to a heterodimeric serine/threonine kinase receptor complex results in the phosphorylation of Mad, a member of the Smad family. Mad 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.)
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)
Component of the Mad-Med transcription factor complex involved in the dpp/BMP TGF-beta signal transduction pathway; not essential but enhances or modifies the signal (PubMed:15296719, PubMed:16109720, PubMed:7705627, PubMed:8702167, PubMed:9502724, PubMed:9694800). Functions downstream of dpp and its receptors (PubMed:9502722, PubMed:9694800). Associates with the R-SMAD Mad upon dpp-signaling and translocates to the nucleus (PubMed:9502724, PubMed:9502733). Can also associate with the activin-like pathway R-SMAD Smox (PubMed:16951053). Binds GC-rich enhancer sites with consensus sequence 5'-GTCTG-3' in the enhancer region of dpp-responsive genes (PubMed:15296719, PubMed:16109720, PubMed:9694800). Mad-Med acts as a transcriptional activator but recruits shn/Shnurri to enhance its activity or switch to transcriptional repression (PubMed:16109720). Regulates dpp-responsive genes by directly activating their expression or by repressing the transcriptional repressor brk/brinker (PubMed:12705870). Also involved in repression of other key developmental genes including bam and gsb (PubMed:15296719). Required for dpp-induced expression of tin/tinman during embryogenesis (PubMed:9694800). Required for dpp-signaling during dorsoventral patterning of the embryo (PubMed:9502724, PubMed:9502733). Maternal contribution is essential for dorsoventral patterning in embryos (PubMed:9502722). Involved in dpp/BMP signaling during development of imaginal discs (PubMed:9502733). ACTIVITY REGULATION: DNA binding of the Mad-Med complex, specifically of the enhancer region of the brk gene, is activated by dpp-signaling (PubMed:15296719). Dpp-signaling induces phosphorylation of Mad by dpp-specific receptor serine/threonine kinase complexes, which in turn induces interaction of Mad with Med and nuclear localization of the Mad-Med complex, where it can bind the enhancer regions of dpp-responsive genes (PubMed:9502724).
(UniProt, O62609)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
l(3)SG70
Homozygous larvae contain rudimentary imaginal discs, but disc primordia do not grow during larval development; testes and ovaries smaller than normal, and cell number in central nervous system reduced. Mutant gonads do not survive metamorphosis when implanted into wild-type larvae. Homozygous cuticular clones appear to develop normally, but with reduced frequency and size compared to control clones. Mutant larvae support growth of implanted wild-type discs. Normal gene product postulated to be required for cell proliferation; survival of somatic epidermal clones attributed to perdurance. Larval ganglion mitoses exhibit weak effect on chromosome condensation as well as chromosome breakage (Gatti and Baker, 1989, Genes Dev. 3: 438-53); salivary chromosomes appear normal.
Summary (Interactive Fly)

Smad family member - associates with Smad1 in response to Dpp or with Smad2 (Smox) in response to Activin ligands - dominant negative Smad4 blocks both BMP and activin responses - developmental roles include dorsal-ventral patterning, patterning and proliferation of the wing disc and gene expression in the mushroom body of the larval brain

Gene Model and Products
Number of Transcripts
3
Number of Unique Polypeptides
2

Please see the JBrowse view of Dmel\Med 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 O62609)

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

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

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Gene model reviewed during 5.48

Gene model reviewed during 5.46

Gene model reviewed during 5.53

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Med-RA
FBtr0085815
NM_079871
3250
771
Med-RB
FBtr0085816
NM_170559
3028
697
Med-RC
FBtr0337030
NM_170560
3425
771
Additional Transcript Data and Comments
Reported size (kB)

2.8 (longest cDNA)

(Das et al., 1998)

3.1-3.3 (northern blot)

(Hudson et al., 1998)

3.3 (longest cDNA)

(Wisotzkey et al., 1998)

3.03 (longest cDNA)

(Xu et al., 1998)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
Med-PA
FBpp0085176
81.6
771
7.60
O62609
NP_524610
AAF57172
Med-PB
FBpp0085177
73.7
697
7.45
Q8IMG6
NP_733438
AAN14277
Med-PC
FBpp0307959
81.6
771
7.60
O62609
NP_733439
AAN14278
Polypeptides with Identical Sequences

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

771 aa isoforms: Med-PA, Med-PC
Additional Polypeptide Data and Comments
Reported size (kDa)

771 (aa)

(Das et al., 1998, Hudson et al., 1998, Wisotzkey et al., 1998)

697 (aa)

(Xu et al., 1998)
Comments
External Data
Subunit Structure (UniProtKB)

Component of a DNA binding Mad-Med complex composed of two copies of Mad and one copy of Med (PubMed:16109720, PubMed:9502724). May form heterodimers composed of one copy of Mad and one copy of Med when associated with certain gene enhancer regions such as the enhancer for brk (PubMed:15296719). Interacts (via C-terminus) with Mad (when phosphorylated at the C-terminus) (PubMed:16109720, PubMed:16951053, PubMed:9502724). The Mad-Med complex associates with shn/Shnurri; recruitment of shn to DNA enhancer elements results in transcriptional repression (PubMed:15296719, PubMed:16109720). Interacts with Smox/SMAD2 (when phosphorylated on the C-terminal SSXS motif); the interaction is dependent on a receptor serine/threonine kinase complex, due to phosphorylation of Smox by the receptor complex (PubMed:10320478, PubMed:16951053). Heterodimerizes with the R-SMAD Smox/SMAD2 upon activation of the activin pathway and translocates to the nucleus, where it transcriptionally activates activin-responsive genes (PubMed:10320478, PubMed:9887103). Interacts with Snoo; the interaction reduces binding of Med to the dpp/BMP-like pathway R-SMAD Mad and promotes binding to the activin-like pathway R-SMAD Smox acting like a pathway switch (PubMed:16951053). Interacts (via N-terminus) with lwr; the interaction is direct (PubMed:18794353).

(UniProt, O62609)
Post Translational Modification

Sumoylated on Lys-141, Lys-185 and Lys-248 by the SUMO-conjugating enzyme lwr/Ubc9 (PubMed:18794353). Sumoylation occurs in the nucleus and promotes nuclear export (PubMed:18794353). Association with phodphorylated Mad prevents sumoylation (PubMed:18794353).

Probably ubiquitinated on Lys-738 (Probable). Deubiquitinated by the ubiquitin hydrolase faf (PubMed:22745309).

(UniProt, O62609)
Domain

The MH1 domain mediates DNA binding (PubMed:15296719, PubMed:9694800). DNA interaction is mediated by a conserved beta-hairpin DNA binding motif (PubMed:16109720).

The MH2 domain has an inhibitory effect on DNA binding.

The C-terminal domain involved in interaction with Mad is required for nuclear localization.

(UniProt, O62609)
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\Med using the Feature Mapper tool.

External Data
Crossreferences
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).

-0.87

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4
(Wisotzkey et al., 1998)
embryonic stage 4 -- 6
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 7
organism | anterior
(Wisotzkey et al., 1998)
mesoderm
(Wisotzkey et al., 1998)
embryonic stage 7 -- 8
organism | ubiquitous
(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)
dorsal ectoderm
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
embryonic stage 11 -- 12
central brain primordium
(Fisher et al., 2012)
dorsal epidermis primordium
(Fisher et al., 2012)
foregut primordium
(Fisher et al., 2012)
hindgut proper primordium
(Fisher et al., 2012)
organism | faint | ubiquitous
(Fisher et al., 2012)
ventral epidermis primordium
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic brain
(Fisher et al., 2012)
embryonic foregut
(Fisher et al., 2012)
embryonic hindgut
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
organism | faint | ubiquitous
(Fisher et al., 2012)
larval ventral nerve cord
(Fisher et al., 2012)
embryonic stage 14
endoderm
(Wisotzkey et al., 1998)
ectoderm
(Wisotzkey et al., 1998)
embryonic stage 14 -- 17
presumptive embryonic/larval central nervous system | restricted
(Wisotzkey et al., 1998)
embryonic stage 16
embryonic midgut constriction 2
(Wisotzkey et al., 1998)
Additional Descriptive Data

Med transcript expression is widespread in the stage 4 embryo. At stage 7, expression is detected in the mesoderm and in the anterior embryo. At stage 14, expression is detected in the ectoderm, endoderm, and the central nervous system. At stage 16, the CNS, the second midgut constriction, as well as other tissues, accumulate Med transcript.

(Wisotzkey et al., 1998)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
oogenesis stage S9 -- S10
centripetal follicle cell
(Muzzopappa and Wappner, 2005)
oogenesis stage S10
follicle cell
(Muzzopappa and Wappner, 2005)
Additional Descriptive Data

Med protein is localized at oogenesis stage S9 to anterior follicle cells that migrate towards the oocyte, and at stage S10 to all follicle cells, with stronger expression observed in stretch cells.

(Muzzopappa and Wappner, 2005)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytoplasm
inferred from high throughput direct assay
(Sarov et al., 2016)
located_in cytosol
inferred from direct assay
(Das et al., 1998, Wisotzkey et al., 1998)
part_of heteromeric SMAD protein complex
inferred from physical interaction with FLYBASE:Smox; FB:FBgn0025800
(Brummel et al., 1999, Das et al., 1999)
inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648
(Kamiya et al., 2008, Wisotzkey et al., 1998)
inferred from direct assay
(Inoue et al., 1998)
located_in nucleus
inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648
(Wisotzkey et al., 1998)
inferred from direct assay
(Das et al., 1998)
part_of RNA polymerase II transcription repressor complex
inferred from physical interaction with FLYBASE:shn; FB:FBgn0003396,FLYBASE:Mad; FB:FBgn0011648
(Yao et al., 2006, Muller et al., 2003)
part_of transcription regulator complex
inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648
(Weiss et al., 2010)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Med 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
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
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 44 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 25 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Med
Transgenic constructs containing regulatory region of Med
Aberrations (Deficiencies and Duplications) ( 11 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal (with Df(3R)tll-e)
lethal (with Med1)
lethal (with Med5)
lethal (with Med13)
lethal (with Medadro)
lethal | maternal effect | recessive
lethal | recessive
lethal - all die before end of larval stage | recessive
lethal - all die before end of pupal stage | recessive
lethal - all die during embryonic stage | germline clone | recessive | rescuable maternal effect
lethal - all die during embryonic stage | maternal effect
lethal - all die during embryonic stage | maternal effect | recessive
lethal - all die during P-stage, with Scer\GAL4Mef2.PR
lethal - all die during P-stage | recessive
viable
viable, with Scer\GAL4tin.CΔ4
Other Phenotypes
Allele
abnormal cell growth | somatic clone
abnormal mitotic cell cycle
abnormal neuroanatomy
abnormal neuroanatomy (with Df(3R)Kpn-A)
abnormal neuroanatomy (with Med3)
abnormal neuroanatomy (with MedC246)
abnormal neuroanatomy | third instar larval stage, with Scer\GAL4ppk.PU
abnormal neurophysiology
abnormal neurophysiology | adult stage
abnormal size
abnormal size | adult stage, with Scer\GAL4A9
abnormal size | adult stage, with Scer\GAL4dsx.KI
abnormal size | adult stage, with Scer\GAL4esg.PU
abnormal size | larval stage
decreased body size | larval stage (with Df(3R)Kpn-A)
decreased cell number
decreased cell number | oogenesis | somatic clone
decreased occurrence of cell division | larval stage
decreased occurrence of cell division | oogenesis
some die during embryonic stage | maternal effect
some die during embryonic stage | recessive | maternal effect | recessive
some die during embryonic stage | recessive | rescuable maternal effect
some die during larval stage | recessive
some die during pupal stage | recessive
visible, with Scer\GAL4A9
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4ptc-559.1
visible | adult stage, with Scer\GAL4A9
visible | adult stage, with Scer\GAL4ey.PU
visible | adult stage, with Scer\GAL4eya.PB
visible | adult stage, with Scer\GAL4GMR.PU
visible | recessive
visible | somatic clone
wild-type
Phenotype manifest in
Allele
adult thorax
amnioserosa
bouton
bouton (with Df(3R)Kpn-A)
bouton (with Med3)
bouton (with MedC246)
cell cycle
chromosome
dense core granule | adult stage, with Scer\GAL4NP0261
egg chorion | maternal effect | somatic clone
egg operculum | maternal effect | somatic clone
embryonic/first instar larval cuticle
embryonic/first instar larval cuticle | maternal effect
embryonic/larval crystal cell
embryonic/larval midgut | embryonic stage
embryonic epidermis
embryonic midgut constriction 2
eye, with Scer\GAL4ey.PU
eye, with Scer\GAL4GMR.PU
eye | decreased number, with Scer\GAL4eya.PB
eye | somatic clone
eye disc | somatic clone
eye disc morphogenetic furrow | somatic clone
eye photoreceptor cell stalk | somatic clone
fat body
female germline stem cell
femur
filzkorper
follicle cell | somatic clone
follicle stem cell | somatic clone
gastric caecum
germarium
imaginal disc
imaginal disc | larval stage
lamina
lamina | somatic clone
larval central nervous system
larval head | dorso-lateral | embryonic stage
larval head | embryonic stage
larval multidendritic class IV neuron | third instar larval stage, with Scer\GAL4ppk.PU
larval somatic muscle cell (with Df(3R)Kpn-A)
leg, with Scer\GAL4ptc-559.1
leg | ectopic, with Scer\GAL4ptc-559.1
lobula columnar neuron LC14 | somatic clone
lysosome | adult stage, with Scer\GAL4dsx.KI
male accessory gland secondary cell, with Scer\GAL4dsx.KI
male accessory gland secondary cell, with Scer\GAL4esg.PU
male accessory gland secondary cell, with Scer\GAL4NP0261
male germline stem cell | germline clone
multivesicular body | adult stage, with Scer\GAL4dsx.KI
neuromuscular junction
neuromuscular junction (with Df(3R)Kpn-A)
neuromuscular junction (with Med3)
neuromuscular junction (with MedC246)
nucleus | adult stage, with Scer\GAL4esg.PU
photoreceptor cell R7
photoreceptor cell R8
posterior crossvein
pretarsus
secretory vesicle | adult stage, with Scer\GAL4dsx.KI
synapse
tarsal segment 4
t-bar
tibia
ventral denticle belt
wing, with Scer\GAL4A9
wing, with Scer\GAL4Bx-MS1096
wing, with Scer\GAL4nub-AC-62
wing, with Scer\GAL4ptc-559.1
wing | cell non-autonomous | somatic clone
wing | somatic clone
wing disc | somatic clone
wing hinge | cell autonomous | somatic clone
wing hinge | cell non-autonomous | somatic clone
wing margin | cell non-autonomous | somatic clone
wing vein, with Scer\GAL4nub-AC-62
wing vein | ectopic, with Scer\GAL4A9
wing vein | ectopic, with Scer\GAL4ptc-559.1
wing vein L3, with Scer\GAL4A9
wing vein L4
wing vein L4, with Scer\GAL4A9
wing vein L5, with Scer\GAL4A9
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\SMAD4
12 of 14
Yes
Yes
8  
Hsap\SMAD1
2 of 14
No
No
1  
Hsap\SMAD2
2 of 14
No
No
1  
Hsap\SMAD3
2 of 14
No
No
1  
Hsap\SMAD5
2 of 14
No
No
2  
Hsap\SMAD6
2 of 14
No
No
2  
Hsap\SMAD7
2 of 14
No
No
1  
Hsap\SMAD9
2 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) (8)
Rnor\Smad4
11 of 14
Yes
Yes
Rnor\Smad1
2 of 14
No
No
Rnor\Smad2
2 of 14
No
No
Rnor\Smad3
2 of 14
No
No
Rnor\Smad5
2 of 14
No
No
Rnor\Smad6
2 of 14
No
No
Rnor\Smad7
2 of 14
No
No
Rnor\Smad9
1 of 14
No
No
Mus musculus (laboratory mouse) (8)
Mmus\Smad4
11 of 14
Yes
Yes
Mmus\Smad1
2 of 14
No
No
Mmus\Smad2
2 of 14
No
No
Mmus\Smad3
2 of 14
No
No
Mmus\Smad5
2 of 14
No
No
Mmus\Smad6
2 of 14
No
No
Mmus\Smad7
2 of 14
No
No
Mmus\Smad9
2 of 14
No
No
Xenopus tropicalis (Western clawed frog) (9)
Xtro\smad4
8 of 13
Yes
Yes
Xtro\smad4.2
5 of 13
No
Yes
Xtro\rpl14
1 of 13
No
No
Xtro\smad1
1 of 13
No
No
Xtro\smad2
1 of 13
No
No
Xtro\smad7
1 of 13
No
No
Xtro\smad9
1 of 13
No
No
Xtro\tmem54
1 of 13
No
No
Xtro\tspan15
1 of 13
No
No
Danio rerio (Zebrafish) (14)
Drer\smad4a
13 of 14
Yes
Yes
Drer\smad10a
13 of 14
Yes
Yes
Drer\gb:cr929477
5 of 14
No
Yes
Drer\smad4b
4 of 14
No
Yes
Drer\smad2
2 of 14
No
No
Drer\smad3a
2 of 14
No
No
Drer\smad3b
2 of 14
No
No
Drer\smad5
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\smad10b
2 of 14
No
Yes
Drer\smad1
1 of 14
No
No
Drer\smad9
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (8)
Cele\sma-4
10 of 14
Yes
Yes
Cele\daf-3
2 of 14
No
Yes
Cele\daf-8
2 of 14
No
No
Cele\sma-2
2 of 14
No
No
Cele\sma-3
2 of 14
No
No
Cele\tag-68
2 of 14
No
No
Cele\C34E11.2
1 of 14
No
No
Cele\daf-14
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (4)
Agam\AgaP_AGAP002902
10 of 12
Yes
Yes
Agam\AgaP_AGAP008551
2 of 12
No
No
Agam\AgaP_AGAP009777
2 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:Med. 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)
Dad
4 of 13
Mad
4 of 13
Smox
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 ( 6 )
Potential Models Based on Orthology ( 3 )
Modifiers Based on Experimental Evidence ( 5 )
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?
SMAD4; SMAD family member 4
12 of 14
SMAD FAMILY MEMBER 4; SMAD4
 
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
Med
suppressible
anchor
(Wang et al., 2018)
Ras85D|p53|CycE|Med
suppressible
aurA
(Sekiya et al., 2023)
Ras85D|p53|CycE|Med
suppressible
aurB
(Sekiya et al., 2023)
Med
suppressible
dpp
(Hudson et al., 1996)
Ras85D|p53|CycE|Med
suppressible
Dsor1
(Sekiya et al., 2023)
Med
suppressible
fuss
(Fischer et al., 2012)
Med
enhanceable
Inx3
(Richard et al., 2017)
Ras85D|p53|CycE|Med
suppressible
Rok
(Sekiya et al., 2023)
Apc|p53|Ras85D|Med
suppressible
sc
(Quintero and Bangi, 2023)
Med
suppressible
Smurf
(Podos et al., 2001)
Ras85D|p53|CycE|Med
suppressible
Src42A
(Sekiya et al., 2023)
Ras85D|p53|CycE|Med
suppressible
Wee1
(Sekiya et al., 2023)
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
anchor
suppressible
Med
(Wang et al., 2018)
ctrip
suppressible
Med
(Keyan et al., 2023)
CycE
suppressible
Med|dpp
(Horsfield et al., 1998)
dpp
enhanceable
Med
(Takaesu et al., 2005, Hudson et al., 1998, Raftery et al., 1995)
dpp
suppressible
Med
(Bangi and Wharton, 2006, Wisotzkey et al., 1998)
dpp|faf
suppressible
Med
(Stinchfield et al., 2012)
dpp|lolal
suppressible
Med
(Quijano et al., 2016)
gbb
suppressible
Med
(Bangi and Wharton, 2006)
hiw
suppressible
Med
(McCabe et al., 2004)
Ras85D
enhanceable
Pten|Apc|Med
(Bangi et al., 2016)
Ras85D
enhanceable
Pten|Med|p53
(Bangi et al., 2016)
Ras85D
enhanceable
Apc|Med|p53
(Bangi et al., 2016)
Ras85D
enhanceable
Pten|Apc|Med|p53
(Bangi et al., 2016)
sax
suppressible
Med
(Das et al., 1998)
Scgδ
suppressible
Med
(Goldstein et al., 2011)
Snoo
suppressible
Med
(Barrio et al., 2007, Takaesu et al., 2006)
sog
enhanceable
Med
(Decotto and Ferguson, 2001)
sog
suppressible
Med
(Stinchfield et al., 2012)
spict
suppressible
Med
(Wang et al., 2007)
tkv
suppressible
Med
(Das et al., 1998)
External Data
Subunit Structure (UniProtKB)
Component of a DNA binding Mad-Med complex composed of two copies of Mad and one copy of Med (PubMed:16109720, PubMed:9502724). May form heterodimers composed of one copy of Mad and one copy of Med when associated with certain gene enhancer regions such as the enhancer for brk (PubMed:15296719). Interacts (via C-terminus) with Mad (when phosphorylated at the C-terminus) (PubMed:16109720, PubMed:16951053, PubMed:9502724). The Mad-Med complex associates with shn/Shnurri; recruitment of shn to DNA enhancer elements results in transcriptional repression (PubMed:15296719, PubMed:16109720). Interacts with Smox/SMAD2 (when phosphorylated on the C-terminal SSXS motif); the interaction is dependent on a receptor serine/threonine kinase complex, due to phosphorylation of Smox by the receptor complex (PubMed:10320478, PubMed:16951053). Heterodimerizes with the R-SMAD Smox/SMAD2 upon activation of the activin pathway and translocates to the nucleus, where it transcriptionally activates activin-responsive genes (PubMed:10320478, PubMed:9887103). Interacts with Snoo; the interaction reduces binding of Med to the dpp/BMP-like pathway R-SMAD Mad and promotes binding to the activin-like pathway R-SMAD Smox acting like a pathway switch (PubMed:16951053). Interacts (via N-terminus) with lwr; the interaction is direct (PubMed:18794353).
(UniProt, O62609 )
Linkouts
Pathways
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3R
Recombination map
3-102
Cytogenetic map
100C7-100D1
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
100C7-100D1
Limits computationally determined from genome sequence between P{EP}pygoEP1076 and P{PZ}ttk02667&P{lacW}ttkj6C7
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
100D-100D
(Hudson, 1997.12.18, Wisotzkey, 1997.10.1, Hudson et al., 1996)
100C4-100D2
(Raftery et al., 1996)
100D-100D
(determined by in situ hybridisation)
(Hudson et al., 1995)
Experimentally Determined Recombination Data
Location

3-102

(FlyBase, 2016)

3-99.1

(Comeron et al., 2012)

3-

(Hudson et al., 1996, Raftery et al., 1996)

3-130.2

(Gatti and Baker, 1989)

3-106

Left of (cM)
(Hudson et al., 1998)
(Wisotzkey et al., 1998)
Right of (cM)
(Decotto and Ferguson, 2001, Wisotzkey et al., 1998)
(Hudson et al., 1998)
Notes

Also mapped to 3-49.3 based on mapping of l(3)SG36, in error.

Stocks and Reagents
Stocks (23)
Genomic Clones (9)
 

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

cDNA Clones (71)
 

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
    Antibody Information
    Laboratory Generated Antibodies
     
    Commercially Available Antibodies
     
    Cell Line Information
    Publicly Available Cell Lines
     
      Other Stable Cell Lines
       
        Other Comments

        DNA-protein interactions: genome-wide binding profile assayed for Med protein in 2-3 hr embryos; see BDTNP1_TFBS_Med collection report.

        (MacArthur et al., 2009)

        RNAi generated by PCR using primers directed to this gene causes a cell growth and viability phenotype when assayed in Kc167 and S2R+ cells.

        (Boutros 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)

        The brk silencer serves as a direct target for a protein complex consisting of Mad/Med and shn.

        (Muller et al., 2003)

        Overexpression of Med or Hsap\MADH4 in the wing and leg causes a similar phenotype. The leg phenotype caused by overexpression of Med or Hsap\MADH4 is similar to that caused by overexpression of Hsap\MADH6 or Hsap\MADH7. Overexpression of Med, Smox, Hsap\MADH2 or Hsap\MADH4 causes a similar phenotype in the wing.

        (Marquez et al., 2001)

        Hsap\MADH1 and Hsap\MADH4, as well as Mad and Med, can stimulate dpp signalling in limb development. Hsap\MADH2, Hsap\MADH4 and Med stimulate activin-β- rather than dpp-mediated cell proliferation. Med can signal for both TGF-beta families - Dpp/BMP signalling Smads and TGF-beta/Activin Smads.

        (Marquez et al., 2001)

        One of five genes identified as encoding downstream components of the dpp signalling cascade which is necessary for blocking salivary gland gene activation by Scr in the dorsal region of parasegment 2. Med function is required to block salivary gland formation in dorsal regions of PS2.

        (Henderson et al., 1999)

        Loss of function alleles of tkv, put, Mad, Med and shn suppress the CycEJP mutant eye phenotype in combination with dppd-ho.

        (Horsfield et al., 1998)

        Med is required for both embryonic and imaginal disc patterning. Med may have two independently mutable functions in patterning the embryonic ectoderm. Med acts downstream of tkv.

        (Hudson et al., 1998)

        Complete elimination of maternal and zygotic Med activity in the early embryo produces a ventralised phenotype identical to that of null dpp mutants, suggesting that Med is required for all dpp-dependent signaling in embryonic dorsal-ventral patterning.

        (Hudson et al., 1998)

        Mutants show no interaction with Df(2R)Pcl11B or Df(3L)66C-G28.

        (Nicholls and Gelbart, 1998)

        Med functions in dpp signaling. Med protein is localised in the cytoplasm, is not regulated by phosphorylation, and requires physical association with Mad protein for nuclear translocation. Mad mediated nuclear translocation is essential for Med function.

        (Wisotzkey et al., 1998)

        A divergent member of the dwarfin family.

        (Cho et al., 1996)

        Dominant enhancer of zen.

        (Hudson et al., 1995)

        Med gene was identified in screens for mutations that decrease apparent activity of the dpp signal in the embryonic ectoderm.

        (Raftery et al., 1995)
        Relationship to Other Genes
        Source for database merge of

        Source for merge of: Med demented

        (Padgett et al., 1997)

        Source for merge of: Med E(zen)3

        (Ferguson, 1998.8.7)

        Source for merge of: Med anon- EST:Posey121

        (Bayraktaroglu, 2000.11.2)
        Additional comments
        Nomenclature History
        Source for database identify of

        Source for identity of: Med CG1775

        (Zhao, 1999.11)
        Nomenclature comments
        Etymology
        Synonyms and Secondary IDs (24)
        Reported As
        Symbol Synonym
        3R16
        (Bellen, 2020.5.15)
        CG1775
        (Neuert et al., 2017, Ni et al., 2011.7.19, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Perkins et al., 2009, Molnar et al., 2006)
        DMT
        (Padgett et al., 1997)
        E(zen)3
        MED
        (Blatti et al., 2015, Restrepo et al., 2014, Slattery et al., 2014, Cheng et al., 2013, MacArthur et al., 2009)
        Med
        (Al Asafen et al., 2024, Eslahi et al., 2024, Fukuda et al., 2024, Jang et al., 2024, Barbaste et al., 2023, Bastin and Eleftherianos, 2023, Lassetter et al., 2023, Liu et al., 2023, Quintero and Bangi, 2023, Sekiya et al., 2023, Bressan and Araujo, 2022, Ding et al., 2022, Jiang et al., 2022, Montanari et al., 2022, Rass et al., 2022, Vicidomini and Serpe, 2022, Zipper et al., 2022, Bilder et al., 2021, Fabian et al., 2021, Finger et al., 2021, Kim and O'Connor, 2021, Li et al., 2021, Sood et al., 2021, Tiwari and Mandal, 2021, Bellen, 2020.5.15, Berndt et al., 2020, Gou et al., 2020, Hegde et al., 2020, Port et al., 2020, Robinson et al., 2020, Zhou and Boutros, 2020, Drummond-Barbosa, 2019, Leiblich et al., 2019, Li et al., 2019, Meltzer et al., 2019, Shokri et al., 2019, Vuilleumier et al., 2019, Ahmed-de-Prado and Baonza, 2018, Bischof et al., 2018, Kwon et al., 2018, Liao et al., 2018, Li et al., 2018, Powers and Srivastava, 2018, Setiawan et al., 2018, Song et al., 2018, Wang et al., 2018, Follansbee et al., 2017, Houtz et al., 2017, Jeibmann et al., 2017, Neuert et al., 2017, Transgenic RNAi Project members, 2017-, Altenhein et al., 2016, Mbodj et al., 2016, Quijano et al., 2016, Sarov et al., 2016, Ayyaz et al., 2015, Bivik et al., 2015, Chen et al., 2015, Newton et al., 2015, Schertel et al., 2015, Zhou et al., 2015, Deshpande et al., 2014, Esteves et al., 2014, Jiang and Singh, 2014, Oh et al., 2014, Pichaud, 2014, Beckwith et al., 2013, Curtis et al., 2013, Hevia and de Celis, 2013, Li and Gilmour, 2013, Saunders et al., 2013, Webber et al., 2013, Fischer et al., 2012, Holmqvist et al., 2012, Kvon et al., 2012, Smith et al., 2012, Spokony and White, 2012.11.14, Stinchfield et al., 2012, Takaesu et al., 2012, Harris and Ashe, 2011, Li et al., 2011, Oh and Irvine, 2011, Quijano et al., 2011, Toku et al., 2011, Ball et al., 2010, Blanco et al., 2010, Chen and Xu, 2010, Dworkin et al., 2009, Lembong et al., 2009, Liu et al., 2009, Casas-Tinto et al., 2008, Christoforou et al., 2008, Miles et al., 2008, Ng, 2008, Pinto et al., 2008, Yakoby et al., 2008, Yao et al., 2008, Beltran et al., 2007, Gao and Laughon, 2007, Li et al., 2007, Shravage et al., 2007, Walsh and Carroll, 2007, Wang et al., 2007, Bangi and Wharton, 2006, Bernardi et al., 2006, Davidson and Erwin, 2006, Gao and Laughon, 2006, Molnar et al., 2006, Sotillos and de Celis, 2006, Takaesu et al., 2006, Yao et al., 2006, Gao et al., 2005, Kirilly et al., 2005, Marques, 2005, Xie et al., 2005, Yamashita et al., 2005, Pyrowolakis et al., 2004, Gim et al., 2001, Kyoda et al., 2000, Pickeral et al., 2000)
        Medea
        (Matsuda and Affolter, 2023, Connacher and Goldstrohm, 2021, Li et al., 2020, Pan et al., 2020, Romero-Pozuelo et al., 2017, Niwa and Niwa, 2016, Urrutia et al., 2016, Van Bortle et al., 2015, Yamanaka et al., 2013, Raftery and Umulis, 2012)
        SMAD4
        (Takaesu et al., 2005)
        Smad4
        (Fu et al., 2024, van Soldt and Cardoso, 2020, Chen and Xu, 2010)
        anon-EST:Posey121
        dSmad4
        (Denton et al., 2020, Bangi et al., 2016, Xu et al., 1998)
        demented
        l(3)11m-254
        (Castellanos et al., 2008, Gatti and Baker, 1989)
        l(3)12m-137
        (Gatti and Baker, 1989)
        l(3)SG36
        l(3)SG70
        (Cohen, 1993, Gatti and Baker, 1989)
        l(3)XIIm137
        med
        (Ell et al., 2024, Deng et al., 2022, Saha et al., 2022, Mahmud et al., 2020, Lu et al., 2019, Whittle and Extavour, 2019, Kang et al., 2018, Wells et al., 2017, Moulton and Letsou, 2016, Ayyaz et al., 2015, Baëza et al., 2015, Kang et al., 2014, Curtis et al., 2013, Marinho et al., 2013, Nègre et al., 2011, The modENCODE Consortium, 2010, The modENCODE Consortium, 2010, Jiang et al., 2008, Ng, 2008, Wang et al., 2008, Kalinovsky and Scheiffele, 2004, Luschnig et al., 2004, Luschnig et al., 2004, McCabe et al., 2002, Baonza and Freeman, 2001, Dequier et al., 2001, Reiter et al., 2001, Horsfield et al., 1998)
        medea
        (Garaulet et al., 2016, Ayyaz et al., 2015, Eaton and Davis, 2005, Deng and Lin, 2001, Kockel et al., 2001, Affolter, 2000, Chen and Fishman, 2000, Potter et al., 2000, Henderson and Andrew, 1998, Haerry et al., 1996)
        smad4
        (Yamamura et al., 2021, Ayyaz et al., 2015)
        Name Synonyms
        Demented
        (Padgett et al., 1997)
        Medea
        (Kinterová et al., 2022, Rass et al., 2022, Lee et al., 2021, Arnés et al., 2020, Li et al., 2020, Schloop et al., 2020, Jordán-Álvarez et al., 2017, Upadhyay et al., 2017, Grill et al., 2016, Bier and De Robertis, 2015, Newton et al., 2015, Li et al., 2014, Mannervik, 2014, Wisotzkey et al., 2014, Zhang et al., 2014, Bausek, 2013, Doumpas et al., 2013, Holmqvist et al., 2012, Sun et al., 2012, Eivers et al., 2011, Eliazer and Buszczak, 2011, Goldstein et al., 2011, Hamaratoglu et al., 2011, Harris and Ashe, 2011, Ball et al., 2010, Bryk et al., 2010, Chen and Xu, 2010, Quijano et al., 2010, Sander et al., 2010, Wagner et al., 2010, Weiss et al., 2010, Lembong et al., 2009, MacArthur et al., 2009, Campbell and Moser, 2008, Jiang et al., 2008, Kamiya et al., 2008, Miles et al., 2008, Miles et al., 2008, Ng, 2008, Padgett, 2008.6.25, Pinto et al., 2008, Warrior et al., 2008, Yakoby et al., 2008, Yao et al., 2008, Barrio et al., 2007, Fuller and Spradling, 2007, Gao and Laughon, 2007, Lilja et al., 2007, Newfeld et al., 2007, Pistillo and Desplan, 2007, Pollard et al., 2007, Potthoff and Olson, 2007, Walsh and Carroll, 2007, Wang et al., 2007, Bernardi et al., 2006, Gao and Laughon, 2006, Li and Li, 2006, Lin et al., 2006, Marques, 2005, Stathopoulos and Levine, 2005, Yamashita et al., 2005, Stefancsik and Sarkar, 2003, Gim et al., 2001, Evangelista et al., 1998, Padgett, 1998.1.7, Padgett et al., 1998, Sutherland and Raftery, 1998, Takase, 1998.4.3, Wisotzkey et al., 1997, Xu, 1997.8.16, Yin, 1997.8.16)
        Media
        (Akasaka et al., 2006)
        demented
        (Cho et al., 1996)
        lethal(3)SG36
        lethal(3)SG70
        medea
        (Sardi et al., 2021, Richard et al., 2017, Luo et al., 2015, Richard and Hoch, 2015, Marinho et al., 2013, Fischer et al., 2012, Nagel et al., 2012, Wang et al., 2007, Nakayama et al., 2006, Eaton and Davis, 2005)
        Secondary FlyBase IDs
        • FBgn0011655
        • FBgn0002479
        • FBgn0002509
        • FBgn0013946
        • FBgn0027760
        • FBgn0015927
        Datasets (2)
        Study focus (2)
        Experimental Role
        Project
        Project Type
        Title
        • bait_protein
        BDTNP_TFBS
        ChIP characterization of transcription factor genome binding, Berkeley Drosophila Transcription Factor Network Project.
        • bait_protein
        modENCODE_regulation_TFs
        Genome-wide localization of transcription factors by ChIP-chip and ChIP-Seq.
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 70 )
        Sequence Crossreferences
        NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
        GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
        GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
        RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
        UniProt/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.
        DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
        EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
        FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
        InterPro - A database of protein families, domains and functional sites
        KEGG Genes - Molecular building blocks of life in the genomic space.
        MARRVEL_MODEL - MARRVEL (model organism gene)
        Linkouts
        FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
        Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
        iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
        Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
        KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
        Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
        References (457)