FB2020_01, released Feb 12, 2020

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
FBgn0011655
Gene Model Status
Current
Stock Availability
18 publicly available
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]
Other Summaries
Also Known As

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

Key Links
Genomic Location
Cytogenetic map
100C7-100D1
Sequence location
3R:31,611,046..31,615,375 [+]
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
GO Summary Ribbons
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
DNA-binding transcription activator activity, RNA polymerase II-specific; DNA-binding transcription repressor activity, RNA polymerase II-specific; protein binding; RNA polymerase II regulatory region sequence-specific DNA binding
Predictions / Assertions
-
Summaries
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.)
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 GBrowse view of Dmel\Med or 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
Comments on Gene Model

Gene model reviewed during 5.48

Gene model reviewed during 5.46

Gene model reviewed during 5.53

Sequence Ontology: Class of Gene
nuclear_gene
protein_coding_gene
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
RefSeq ID
GenBank
Med-PA
FBpp0085176
81.6
771
7.60
NP_524610
AAF57172
Med-PB
FBpp0085177
73.7
697
7.45
NP_733438
AAN14277
Med-PC
FBpp0307959
81.6
771
7.60
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
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
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (30 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
DNA-binding transcription activator activity, RNA polymerase II-specific
inferred from direct assay
(Weiss et al., 2010)
DNA-binding transcription repressor activity, RNA polymerase II-specific
inferred from direct assay
(Yao et al., 2006, Muller et al., 2003)
protein binding
inferred from physical interaction with UniProtKB:Q9NB71
(assigned by UniProt )
(McCabe et al., 2004)
RNA polymerase II regulatory region sequence-specific DNA binding
inferred from direct assay
(Weiss et al., 2010)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (20 terms)
Terms Based on Experimental Evidence (17 terms)
CV Term
Evidence
References
activin receptor signaling pathway
inferred from direct assay
(Brummel et al., 1999, Das et al., 1999)
BMP signaling pathway
inferred from genetic interaction with FLYBASE:dpp; FB:FBgn0000490
(Hudson et al., 1998, Wisotzkey et al., 1998, Raftery et al., 1995)
inferred from direct assay
(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, FLYBASE:sax; FB:FBgn0003317
(Das et al., 1998)
inferred from mutant phenotype
(McCabe et al., 2004, Hudson et al., 1998)
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)
compound eye morphogenesis
inferred from mutant phenotype
(Das et al., 1998)
dorsal/ventral axis specification
inferred from mutant phenotype
(Hudson et al., 1998)
dorsal/ventral pattern formation
inferred from mutant phenotype
(Das et al., 1998, Wisotzkey et al., 1998)
germ-line stem cell division
inferred from mutant phenotype
(Xie and Spradling, 1998)
germ-line stem cell population maintenance
inferred from mutant phenotype
(Xie and Spradling, 1998)
imaginal disc-derived leg morphogenesis
inferred from mutant phenotype
(Marquez et al., 2001)
imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Marquez et al., 2001)
negative regulation of transcription by RNA polymerase II
inferred from direct assay
(Yao et al., 2006, Muller et al., 2003)
neuron development
inferred from mutant phenotype
(Zheng et al., 2006)
ovarian follicle cell development
inferred from mutant phenotype
(Kirilly et al., 2005)
positive regulation of synaptic growth at neuromuscular junction
inferred from mutant phenotype
(McCabe et al., 2004)
positive regulation of transcription by RNA polymerase II
inferred from direct assay
(Weiss et al., 2010, Dai et al., 2000, Das et al., 1999)
retinal ganglion cell axon guidance
inferred from mutant phenotype
(Yoshida et al., 2005)
SMAD protein signal transduction
inferred from direct assay
(Das et al., 1998)
somatic stem cell population maintenance
inferred from mutant phenotype
(Kirilly et al., 2005)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
heart development
traceable author statement
(Cripps and Olson, 2002, Chen and Fishman, 2000)
negative regulation of salivary gland boundary specification
traceable author statement
(Bradley et al., 2001)
transforming growth factor beta receptor signaling pathway
inferred from electronic annotation with InterPro:IPR013019, InterPro:IPR036578
(assigned by InterPro )
(FlyBase Curators et al., 2004-)
Cellular Component (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
cytoplasm
inferred from high throughput direct assay
(Sarov et al., 2016)
cytosol
inferred from direct assay
(Das et al., 1998, Wisotzkey et al., 1998)
heteromeric SMAD protein complex
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)
inferred from physical interaction with FLYBASE:Smox; FB:FBgn0025800
(Brummel et al., 1999, Das et al., 1999)
nucleus
inferred from direct assay
(Das et al., 1998)
inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648
(Wisotzkey et al., 1998)
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)
transcription factor complex
inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648
(Weiss et al., 2010)
Terms Based on Predictions or Assertions (0 terms)
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 7
organism | anterior
(Wisotzkey et al., 1998)
mesoderm
(Wisotzkey et al., 1998)
embryonic stage 16
embryonic midgut constriction 2
(Wisotzkey et al., 1998)
embryonic stage 14 -- 17
presumptive embryonic/larval central nervous system | restricted
(Wisotzkey et al., 1998)
embryonic stage 14
endoderm
(Wisotzkey et al., 1998)
ectoderm
(Wisotzkey et al., 1998)
embryonic stage 4
(Wisotzkey et al., 1998)
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
organism | ubiquitous
(Fisher et al., 2012)
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)
ventral nerve cord
(Fisher et al., 2012)
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
centripetally migrating 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
cytoplasm
inferred from high throughput direct assay
(Sarov et al., 2016)
cytosol
inferred from direct assay
(Das et al., 1998, Wisotzkey et al., 1998)
heteromeric SMAD protein complex
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)
inferred from physical interaction with FLYBASE:Smox; FB:FBgn0025800
(Brummel et al., 1999, Das et al., 1999)
nucleus
inferred from direct assay
(Das et al., 1998)
inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648
(Wisotzkey et al., 1998)
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)
transcription factor complex
inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648
(Weiss et al., 2010)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Med in GBrowse 2
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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
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, and Transgenic Constructs
Classical and Insertion Alleles ( 40 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 20 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Med
Transgenic constructs containing regulatory region of Med
Deletions and Duplications ( 11 )
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 embryonic stage | germline clone | recessive | rescuable maternal effect
lethal - all die before end of embryonic stage | maternal effect
lethal - all die before end of embryonic stage | maternal effect | recessive
lethal - all die before end of larval stage | recessive
lethal - all die before end of pupal stage | recessive
lethal - all die during P-stage, with Scer\GAL4Mef2.PR
viable
viable, with Scer\GAL4tin.CΔ4
Other Phenotypes
Allele
cell growth defective | somatic clone
decreased cell number
decreased cell number | oogenesis | oogenesis | somatic clone
decreased occurrence of cell division | larval stage
decreased occurrence of cell division | oogenesis | oogenesis
mitotic cell cycle defective
neuroanatomy defective
neuroanatomy defective (with Df(3R)Kpn-A)
neuroanatomy defective (with Med3)
neuroanatomy defective (with MedC246)
neuroanatomy defective | third instar larval stage, with Scer\GAL4ppk.PU
neurophysiology defective
size defective
size defective | adult stage, with Scer\GAL4A9
size defective | adult stage, with Scer\GAL4dsx.KI
size defective | adult stage, with Scer\GAL4esg.PU
size defective | larval stage
small body | larval stage (with Df(3R)Kpn-A)
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 | 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
chorion | maternal effect | somatic clone
chromosome
crystal cell
dense core granule | adult stage, with Scer\GAL4NP0261
denticle belt
dorsal cluster neuron | somatic clone
egg operculum | maternal effect | somatic clone
embryonic/first instar larval cuticle
embryonic/first instar larval cuticle | maternal effect
embryonic/larval central nervous system
embryonic/larval midgut | embryonic stage
embryonic/larval somatic muscle (with Df(3R)Kpn-A)
embryonic epidermis
embryonic midgut constriction 2
eye | somatic clone
eye disc | 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 | somatic clone
larval head | dorso-lateral | embryonic stage
larval head | embryonic stage
larval multidendritic class IV neuron | third instar larval stage, with Scer\GAL4ppk.PU
leg, with Scer\GAL4ptc-559.1
leg | ectopic, with Scer\GAL4ptc-559.1
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
morphogenetic furrow | somatic 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
posterior crossvein
pretarsus
secretory vesicle | adult stage, with Scer\GAL4dsx.KI
synapse
tarsal segment 4
t-bar
tibia
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
Download All OrthoDB Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (8)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Hsap\SMAD4
12 of 15
Yes
Yes
 
Hsap\SMAD1
1 of 15
No
No
 
Hsap\SMAD2
1 of 15
No
No
 
Hsap\SMAD3
1 of 15
No
No
 
Hsap\SMAD5
1 of 15
No
No
 
Hsap\SMAD6
1 of 15
No
No
 
Hsap\SMAD7
1 of 15
No
No
 
Hsap\SMAD9
1 of 15
No
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (8)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Mmus\Smad4
13 of 15
Yes
Yes
Mmus\Smad1
1 of 15
No
No
Mmus\Smad2
1 of 15
No
No
Mmus\Smad3
1 of 15
No
No
Mmus\Smad5
1 of 15
No
No
Mmus\Smad6
1 of 15
No
No
Mmus\Smad7
1 of 15
No
No
Mmus\Smad9
1 of 15
No
No
Rattus norvegicus (Norway rat) (8)
Rnor\Smad4
8 of 13
Yes
Yes
Rnor\LOC103691556
1 of 13
No
No
Rnor\Smad1
1 of 13
No
No
Rnor\Smad2
1 of 13
No
No
Rnor\Smad3
1 of 13
No
No
Rnor\Smad5
1 of 13
No
No
Rnor\Smad6
1 of 13
No
No
Rnor\Smad7
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (9)
Xtro\smad4.1
8 of 12
Yes
Yes
Xtro\smad4.2
3 of 12
No
Yes
Xtro\smad1
1 of 12
No
No
Xtro\smad2
1 of 12
No
No
Xtro\smad3
1 of 12
No
No
Xtro\smad6.2
1 of 12
No
No
Xtro\smad6
1 of 12
No
No
Xtro\smad7
1 of 12
No
No
Xtro\smad9
1 of 12
No
No
Danio rerio (Zebrafish) (12)
Drer\si:dkey-222b8.1
11 of 15
Yes
Yes
Drer\si:dkey-239n17.4
11 of 15
Yes
Yes
Drer\smad4a
10 of 15
No
Yes
Drer\gb:cr929477
7 of 15
No
Yes
Drer\smad2
1 of 15
No
No
Drer\smad3a
1 of 15
No
No
Drer\smad3b
1 of 15
No
No
Drer\smad5
1 of 15
No
No
Drer\smad6a
1 of 15
No
No
Drer\smad6b
1 of 15
No
No
Drer\smad7
1 of 15
No
No
Drer\smad9
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (7)
Cele\sma-4
8 of 15
Yes
Yes
Cele\daf-3
5 of 15
No
Yes
Cele\daf-8
1 of 15
No
No
Cele\daf-14
1 of 15
No
No
Cele\sma-2
1 of 15
No
No
Cele\sma-3
1 of 15
No
No
Cele\tag-68
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG0919062J )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Dmel\Med
Drosophila suzukii
Spotted wing Drosophila
Dsuz\DS10_00007391
Drosophila simulans
Dsim\GD21590
Drosophila sechellia
Dsec\GM16436
Drosophila erecta
Dere\GG11819
Drosophila yakuba
Dyak\GE10952
Drosophila ananassae
Dana\GF22974
Drosophila pseudoobscura pseudoobscura
Dpse\GA14643
Drosophila persimilis
Dper\GL23637
Drosophila willistoni
Dwil\GK11724
Drosophila virilis
Dvir\GJ23092
Drosophila mojavensis
Dmoj\GI23400
Drosophila grimshawi
Dgri\GH18642
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG0915018A )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Mdoa\17005647
Glossina morsitans
Tsetse fly
Gmor\GMOY010539
Lucilia cuprina
Australian sheep blowfly
Lcup\FF38_00266
Aedes aegypti
Yellow fever mosquito
Aaeg\AAEL013896
Anopheles darlingi
American malaria mosquito
Adar\ADAC007516
Anopheles gambiae
Malaria mosquito
Agam\AGAP002902
Culex quinquefasciatus
Southern house mosquito
Cqui\CPIJ002563
Culex quinquefasciatus
Southern house mosquito
Cqui\CPIJ002562
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W03OZ )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bmor\BGIBMGA010110
Danaus plexippus
Monarch butterfly
Dple\DPOGS207335-PA
Heliconius melpomene
Postman butterfly
Hmel\HMEL002203
Apis florea
Little honeybee
Aflo\3081814724
Apis mellifera
Western honey bee
Amel\GB50071
Bombus impatiens
Common eastern bumble bee
Bimp\BIMP16290
Bombus terrestris
Buff-tailed bumblebee
Bter\15205669
Linepithema humile
Argentine ant
Lhum\LH21571
Megachile rotundata
Alfalfa leafcutting bee
Mrot\MROTU:001a25
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG006495
Dendroctonus ponderosae
Mountain pine beetle
Dpod\YQE_06624
Tribolium castaneum
Red flour beetle
Tcas\Med
Pediculus humanus
Human body louse
Phum\PHUM002670
Cimex lectularius
Bed bug
Clec\CLEC002593
Cimex lectularius
Bed bug
Clec\CLEC002594
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI000365
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI003759
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI005550
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI008450
Zootermopsis nevadensis
Nevada dampwood termite
Znev\Znev_02071
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X03M2 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Smar\SMAR013850
Ixodes scapularis
Black-legged tick
Isca\ISCW007667
Ixodes scapularis
Black-legged tick
Isca\ISCW018468
Stegodyphus mimosarum
African social velvet spider
Smim\KFM60390.1
Tetranychus urticae
Two-spotted spider mite
Turt\tetur10g00400
Daphnia pulex
Water flea
Dpux\DAPPUDRAFT_304115
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G05Z9 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Smad4
Ciona intestinalis
Vase tunicate
Cint\ENSCING00000005559
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (3)
Gene Symbol
Score
Source
Compara
eggNOG
Homologene
Inparanoid
Isobase
OrthoDB
OrthoFinder
orthoMCL
Panther
RoundUp
Alignment
Dad
3 of 10
Mad
3 of 10
Smox
3 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Potential Models Based on Orthology ( 2 )
Human Ortholog
Disease
Evidence
References
SMAD4; SMAD family member 4
model of  pancreatic carcinoma
IEA
(FlyBase, 2019-)
model of  juvenile polyposis syndrome
IEA
(FlyBase, 2019-)
Modifiers Based on Experimental Evidence ( 3 )
Allele
Disease
Interaction
References
Med13
ameliorates  muscular dystrophy
modeled by Scgδ840
(Goldstein et al., 2011)
ameliorates  cardiomyopathy
modeled by Scgδ840
(Goldstein et al., 2011)
Med15
ameliorates  muscular dystrophy
modeled by Scgδ840
(Goldstein et al., 2011)
Med17
ameliorates  muscular dystrophy
modeled by Scgδ840
(Goldstein et al., 2011)
Disease Associations of Human Orthologs (via DIOPT v7.1 and OMIM)
Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
Homo sapiens (Human)
Gene name
Score
OMIM
OMIM Phenotype
DO term
Complementation?
Transgene?
SMAD4; SMAD family member 4
12 of 15
600993
 
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
esyN Network Diagram
esyN Network Key:
Suppression
Enhancement
View in Interactions Browser
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)
Med
suppressible
dpp
(Hudson et al., 1996)
Med
suppressible
fuss
(Fischer et al., 2012)
Med
suppressible
Smurf
(Podos et al., 2001)
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
anchor
suppressible
Med
(Wang et al., 2018)
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
Med|Pten|p53
(Bangi et al., 2016)
Ras85D
enhanceable
Apc|Med|p53
(Bangi et al., 2016)
Ras85D
enhanceable
Apc|Med|Pten
(Bangi et al., 2016)
Ras85D
enhanceable
Apc|Med|Pten|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
suppressible
Med
(Stinchfield et al., 2012)
sog
enhanceable
Med
(Decotto and Ferguson, 2001)
spict
suppressible
Med
(Wang et al., 2007)
tkv
suppressible
Med
(Das et al., 1998)
External Data
Linkouts
BioGRID - A database of protein and genetic interactions.
DroID - A comprehensive database of gene and protein interactions.
InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Signaling Pathways (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.)
Metabolic Pathways
External Data
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3R
Recombination map

3-102

Cytogenetic map
100C7-100D1
Sequence location
3R:31,611,046..31,615,375 [+]
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 (18)
Genomic Clones (9)
 

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

cDNA Clones (73)
 

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

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

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

cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones
Other clones
RNAi and Array Information
Linkouts
DRSC - Results frm RNAi screens
GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
Antibody Information
Laboratory Generated Antibodies
 
Commercially Available Antibodies
 
Other Information
Relationship to Other Genes
Source for database identify of

Source for identity of: Med CG1775

(Zhao, 1999.11)
Source for database merge of

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
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)

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)
Origin and Etymology
Discoverer
Etymology
Identification
External Crossreferences and Linkouts ( 83 )
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/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
Other crossreferences
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
InterPro - A database of protein families, domains and functional sites
KEGG Genes - Molecular building blocks of life in the genomic space.
modMine - A data warehouse for the modENCODE project
SignaLink - A signaling pathway resource with multi-layered regulatory networks.
Linkouts
BioGRID - A database of protein and genetic interactions.
DroID - A comprehensive database of gene and protein interactions.
DRSC - Results frm RNAi screens
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
FlyMine - An integrated database for Drosophila genomics
Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
Synonyms and Secondary IDs (20)
Reported As
Symbol Synonym
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)
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
(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, 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, 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, Li and Gilmour, 2013, Saunders et al., 2013, Webber et al., 2013, Fischer et al., 2012, Holmqvist et al., 2012, Kvon 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
(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
(Chen and Xu, 2010)
anon-EST:Posey121
dSmad4
(Bangi et al., 2016, Xu et al., 1998)
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
(Lu et al., 2019, Whittle and Extavour, 2019, Kang et al., 2018, Wells et al., 2017, Moulton and Letsou, 2016, Ayyaz et al., 2015, Kang et al., 2014, 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
(Ayyaz et al., 2015)
Name Synonyms
Medea
(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, 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)
lethal(3)SG36
lethal(3)SG70
medea
(Richard et al., 2017, 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
  • FBgn0002479
  • FBgn0002509
  • FBgn0013946
  • FBgn0027760
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.
References (383)