FB2019_05, released Oct 15, 2019

Gene: Dmel\Mad

Open Close
General Information
Symbol
Dmel\Mad
Species
D. melanogaster
Name
Mothers against dpp
Annotation Symbol
CG12399
Feature Type
protein_coding_gene
FlyBase ID
FBgn0011648
Gene Model Status
Current
Stock Availability
28 publicly available
Gene Snapshot
Mothers against dpp (Mad) encodes the primary transcription factor that mediates cellular response to the BMP like ligands encoded by dpp, scw and gbb. Upon phosphorylation by either the products of sax or tkv (type I BMP receptors), it forms a complex with the product of Med and translocates to the nucleus where, together with cofactors, it regulates expression of BMP response target genes. [Date last reviewed: 2019-03-14]
Other Summaries
Also Known As
pMad, Mothers Against Decapentaplegic, Smad, Mother against decapentaplegic, dMad
Key Links
Genomic Location
Cytogenetic map
23D3-23D3
Sequence location
2L:3,146,056..3,159,643 [-]
Recombination map
2-8
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)
Belongs to the dwarfin/SMAD family. (P42003)
Protein Signatures (InterPro)
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
DNA-binding transcription activator activity, RNA polymerase II-specific; DNA-binding transcription factor activity, RNA polymerase II-specific; DNA-binding transcription repressor activity, RNA polymerase II-specific; protein binding; RNA polymerase II activating transcription factor binding; RNA polymerase II distal enhancer sequence-specific DNA binding; RNA polymerase II regulatory region DNA binding; RNA polymerase II regulatory region sequence-specific DNA binding; sequence-specific DNA binding; transcription coactivator activity
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.)
Protein Function (UniProtKB)
Required for the function of decapentaplegic. May play an important role in mediating Dpp signaling. Involved in the BMP signaling pathway.
(UniProt, P42003)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
apg: apang
Homozygotes when raised at 19 show occasional absence of one or both claws; veins L4 and L5 interrupted; fertile at 19 but become sterile when shifted to 28; produce embryos with range of germ band abnormalities. Homozygous pupal lethal when raised at 28; pharate adults show defective tarsal development of all six legs; condensed, poorly developed and curved metatarsus and tarsi; duplications in tibial and tarsal segments; claws absent. Temperature sensitive period first instar to early pupa.
Summary (Interactive Fly)
Smad family members convey TGFß signals from their receptors to the nucleus - Upon phosphorylation by either Sax or Tkv (type I BMP receptors), Mad forms a complex with Med and translocates to the nucleus where, together with cofactors, it regulates expression of BMP response target genes.
Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
2

Please see the GBrowse view of Dmel\Mad or the JBrowse view of Dmel\Mad 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 5.45
Gene model reviewed during 5.46
Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated
Sequence Ontology: Class of Gene
nuclear_gene
protein_coding_gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Mad-RA
FBtr0077616
NM_057669
2691
455
Mad-RB
FBtr0332372
NM_001273063
3262
525
Additional Transcript Data and Comments
Reported size (kB)
2.6 (northern blot)
(Sekelsky et al., 1995)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
Mad-PA
FBpp0077302
50.5
455
7.04
NP_477017
AAF51142
Mad-PB
FBpp0304648
57.9
525
7.20
NP_001259992
AGB92529
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)
455 (aa)
(Sekelsky et al., 1995)
Comments
External Data
Subunit Structure (UniProtKB)
Homotrimer (PubMed:19557331). Interacts with MAN1 (PubMed:20036230). Interacts with Sec13 and Nup93-1 (PubMed:20547758).
(UniProt, P42003)
Post Translational Modification
Phosphorylation on Ser-453 and/or Ser-455 is required for interaction with Smurf (PubMed:12754252, PubMed:18327897). Phosphorylation on Ser-25 by key/Nemo promotes export from nucleus and antagonizes BMP signaling (PubMed:17507407). Ubiquitinated by Smurf upon phosphorylation; which promotes proteasomal degradation.
(UniProt, P42003)
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\Mad 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 (42 terms)
Molecular Function (10 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
DNA-binding transcription activator activity, RNA polymerase II-specific
inferred from direct assay
(Weiss et al., 2010, Saller and Bienz, 2001)
DNA-binding transcription factor activity, RNA polymerase II-specific
inferred from direct assay
(Hodar et al., 2014, Kim et al., 1997)
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:Q9V853
(assigned by UniProt )
(Liang et al., 2003, Podos et al., 2001)
inferred from physical interaction with FLYBASE:vis; FB:FBgn0033748
(Hyman et al., 2003)
inferred from physical interaction with UniProtKB:Q02748
(Li and Li, 2006)
RNA polymerase II activating transcription factor binding
inferred from physical interaction with FLYBASE:shn; FB:FBgn0003396
(Dai et al., 2000)
RNA polymerase II distal enhancer sequence-specific DNA binding
inferred from direct assay
(Xu et al., 1998)
RNA polymerase II regulatory region DNA binding
inferred from direct assay
(Muller et al., 2003)
RNA polymerase II regulatory region sequence-specific DNA binding
inferred from direct assay
(Weiss et al., 2010, Saller and Bienz, 2001)
sequence-specific DNA binding
inferred from direct assay
(Lin et al., 2006, Grienenberger et al., 2003)
transcription coactivator activity
inferred from direct assay
(Dai et al., 2000)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (26 terms)
Terms Based on Experimental Evidence (23 terms)
CV Term
Evidence
References
BMP signaling pathway
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:tkv; FB:FBgn0003716
(Das et al., 1998)
inferred from mutant phenotype
(Fuentes-Medel et al., 2012, Gesualdi and Haerry, 2007, McCabe et al., 2004, Minami et al., 1999, Lecuit et al., 1996)
inferred from genetic interaction with FLYBASE:dpp; FB:FBgn0000490
(Wisotzkey et al., 1998, Newfeld et al., 1996, Wiersdorff et al., 1996, Raftery et al., 1995, Sekelsky et al., 1995)
inferred from genetic interaction with FLYBASE:tkv; FB:FBgn0003716
(Wells et al., 2017, Wiersdorff et al., 1996)
inferred from genetic interaction with FLYBASE:wg; FB:FBgn0284084
(Wiersdorff et al., 1996)
compound eye morphogenesis
inferred from mutant phenotype
(Cordero et al., 2007)
inferred from genetic interaction with FLYBASE:dpp; FB:FBgn0000490
(Wiersdorff et al., 1996)
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)
histoblast morphogenesis
inferred from mutant phenotype
(Ninov et al., 2010)
imaginal disc morphogenesis
inferred from mutant phenotype
(Sekelsky et al., 1995)
imaginal disc-derived leg morphogenesis
inferred from mutant phenotype
(Marquez et al., 2001)
imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Dworkin and Gibson, 2006, Lecuit et al., 1996)
imaginal disc-derived wing vein morphogenesis
inferred from mutant phenotype
(Marquez et al., 2001)
negative regulation of G1/S transition of mitotic cell cycle
inferred from mutant phenotype
(Firth and Baker, 2005)
negative regulation of transcription by RNA polymerase II
inferred from direct assay
(Yao et al., 2006, Muller et al., 2003)
inferred from mutant phenotype
(Minami et al., 1999)
ovarian follicle cell development
inferred from mutant phenotype
(Kirilly et al., 2005)
positive regulation of neuromuscular junction development
inferred from mutant phenotype
(Fuentes-Medel et al., 2012)
positive regulation of synaptic growth at neuromuscular junction
inferred from mutant phenotype
(Merino et al., 2009, McCabe et al., 2004)
positive regulation of transcription by RNA polymerase II
inferred from direct assay
(Hodar et al., 2014, Weiss et al., 2010, Saller and Bienz, 2001, Dai et al., 2000)
inferred from expression pattern
(Kim et al., 1997)
inferred from mutant phenotype
(Grienenberger et al., 2003, Xu et al., 1998)
R8 cell fate specification
inferred from mutant phenotype
(Wells et al., 2017)
regulation of cell differentiation
inferred from mutant phenotype
(Mathur et al., 2010)
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)
transforming growth factor beta receptor signaling pathway involved in endodermal cell fate specification
inferred from mutant phenotype
(Guo et al., 2013)
trunk segmentation
inferred from mutant phenotype
(Eivers et al., 2009)
ventral cord development
inferred from high throughput mutant phenotype
(Bivik et al., 2015)
wing disc anterior/posterior pattern formation
inferred from mutant phenotype
(Shen and Dahmann, 2005)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
dorsal closure
non-traceable author statement
(Harden, 2002)
heart development
traceable author statement
(Cripps and Olson, 2002)
negative regulation of salivary gland boundary specification
traceable author statement
(Bradley et al., 2001)
ovarian follicle cell development
traceable author statement
(Dobens and Raftery, 2000)
Cellular Component (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
cytoplasm
inferred from direct assay
(Chen et al., 2006)
cytosol
inferred from direct assay
(Li and Li, 2006, Das et al., 1998, Wisotzkey et al., 1998, Newfeld et al., 1996)
heteromeric SMAD protein complex
inferred from physical interaction with FLYBASE:Med; FB:FBgn0011655
(Kamiya et al., 2008, Wisotzkey et al., 1998)
inferred from direct assay
(Inoue et al., 1998)
nucleus
inferred from direct assay
(Friedman et al., 2013, Chen et al., 2006, Li and Li, 2006, Das et al., 1998, Inoue et al., 1998, Wisotzkey et al., 1998, Kim et al., 1997)
inferred from direct assay
(assigned by UniProt )
(Colozza et al., 2011)
RNA polymerase II transcription repressor complex
inferred from physical interaction with FLYBASE:shn; FB:FBgn0003396, FLYBASE:Med; FB:FBgn0011655
(Yao et al., 2006, Muller et al., 2003)
transcription factor complex
inferred from physical interaction with FLYBASE:Med; FB:FBgn0011655
(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 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
organism
(Fisher et al., 2012)
embryonic stage 11 -- 12
anterior midgut primordium
(Fisher et al., 2012)
garland cell primordium
(Fisher et al., 2012)
posterior midgut primordium
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic/larval garland cell
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage -- adult stage
(Sekelsky et al., 1995)
Additional Descriptive Data
Mad 2.6kb transcripts are detected in all developmental stages but are most abundant in early embryos, pupae, and adult females. A minor slightly smaller transcript is also detected in 0-4hr embryos.
(Sekelsky et al., 1995)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
organism | ubiquitous
(Newfeld et al., 1996)
embryonic stage 5
organism | dorsal

Comment: Expressed in a dorsal midline stripe

(Srinivasan et al., 2002)
embryonic stage 11
amnioserosa

Comment: pMad expression

(Fernández et al., 2007)
organism | ventral

Comment: pMad expression

(Fernández et al., 2007)
embryonic epidermis | dorsal

Comment: pMad expression

(Fernández et al., 2007)
embryonic stage 12 -- 14
embryonic epidermis | striped

Comment: pMad expression

(Fernández et al., 2007)
embryonic stage 13
embryonic dorsal epidermis
(Zahedi et al., 2008)
embryonic stage 13 -- 17
spiracular chamber of embryonic/larval posterior spiracle
(Takaesu et al., 2008)
embryonic stage 14 -- 17
embryonic/larval spiracular branch
(Takaesu et al., 2008)
embryonic/larval dorsal branch
(Takaesu et al., 2008)
embryonic stage 15
neuron | subset of presumptive embryonic/larval central nervous system
(Rawson et al., 2003)
embryonic stage 15 -- 16
embryonic/larval midgut
(Rawson et al., 2003)
embryonic stage 16 -- first instar larval stage
RP2 motor neuron
(Rawson et al., 2003)
RP3 motor neuron
(Rawson et al., 2003)
aCC neuron
(Rawson et al., 2003)
U neuron
(Rawson et al., 2003)
larval stage
apterous-expressing neuron of the lateral cluster of the thorax Ap4
(Baumgardt et al., 2007)
larval CCAP A1-A4 EN abdominal neuron
(Veverytsa and Allan, 2011)
third instar larval stage
germline stem cell
• nucleus

Comment: in primordial germ cell (PGC), at mid-third instar (ML3, 96hr AEL)

(Gilboa and Lehmann, 2004)
embryonic/larval neuromuscular junction | restricted

Comment: phosphorylated Mad (P-Mad)

(Nahm et al., 2010)
morphogenetic furrow

Comment: antibody specific to phosphorylated Mad

(Yuva-Aydemir et al., 2011, Firth et al., 2010)
eye disc anterior to the morphogenetic furrow | restricted

Comment: antibody specific to phosphorylated Mad

(Yuva-Aydemir et al., 2011)
embryonic/larval neuromuscular junction

Comment: antibody specific to phosphorylated Mad (pMad)

(Higashi-Kovtun et al., 2010)
embryonic/larval neuromuscular junction

Comment: antibody specific to phosphorylated Mad (pMad)

(Higashi-Kovtun et al., 2010)
motor neuron of embryonic/larval neuromuscular junction
• nucleus

Comment: antibody specific to phosphorylated Mad (pMad)

(Higashi-Kovtun et al., 2010)
ventral nerve cord
• nucleus

Comment: antibody specific to phosphorylated Mad (pMad)

(Higashi-Kovtun et al., 2010)
eye disc anterior to the morphogenetic furrow
(Firth et al., 2010)
peripodial epithelium of eye disc
(Firth et al., 2010)
antennal disc | ventral
(Firth et al., 2010)
peripodial epithelium of antennal disc
(Firth et al., 2010)
wandering third instar larval stage
anterior-posterior compartment boundary of the wing disc | gradient
(Bornemann et al., 2004)
pupal stage
larval CCAP A1-A4 EN abdominal neuron
(Veverytsa and Allan, 2011)
pupal stage P6
wing vein

Comment: pMad

(Conley et al., 2000)
presumptive wing vein
(Shimmi et al., 2005)
posterior crossvein
(Shimmi et al., 2005)
longitudinal vein
(Shimmi et al., 2005)
pupal stage P6 -- P7
crossvein

Comment: 23-26 hr APF

(Conley et al., 2000)
adult stage | female
female germline stem cell

Comment: high expression level

(Song et al., 2004)
cystoblast

Comment: low expression level

(Song et al., 2004)
adult stage | male
male germline stem cell
(Kawase et al., 2004)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
anterior-posterior compartment boundary of the wing disc

Comment: anti-phosphorylated Mad antibody

(Chauhan et al., 2013)
Additional Descriptive Data
pMad is expressed in CCAP-EN neurons but not in the CCAP-IN neurons.
(Veverytsa and Allan, 2011)
The phosphorylated form of Mad protein is detected in two stripes in the eye disc: one corresponding to the morphogenetic furrow and the other anterior to it. No expression of phosphorylated Mad is detected in glial cells in the eye disc.
(Yuva-Aydemir et al., 2011)
In the eye disc, antibody to phosphorylated Mad stains a broad band of cells anterior to and within the morphogenetic furrow, terminating in a stripe of more intensely labelled cells around columns 3 and 4 at the posterior edge of the furrow. In antennal disc, a broad band of nuclei in the ventral region of the disc is stained. Staining is observed in peripodial cells over the ventral region of the antennal disc and the dorsal region of the eye disc. In some preparations, nuclear pMad was also weakly detected in peripodial epithelium cells directly overlying the morphogenetic furrow.
(Firth et al., 2010)
Phosphorylated Mad protein is expressed in the presynaptic domain of type I boutons at the neuromuscular junction.
(Nahm et al., 2010)
Phosphorylated Mad (pMAd) protein is detected in the spiracular chambers, spiracular branches, and dorsal trunk branches of the tracheal system. Expression is seen in the spiracular chamber form embryonic stage 13 and appears in the spriacular branch and dorsal trunk branches at stage 14 and persists through stage 17.
(Takaesu et al., 2008)
Mad is observed in the dorsal epidermis and the ventrolateral epidermis at embryonic stage 13.
(Zahedi et al., 2008)
Phosphorylated Mad (pMad) is detected in the amnioserosa and in the epidermis around the leading edge at embryonic stage 11. A ventral stripe is also seen. From stage 12 on, pMad levels decline in the amnioserosa but persist in epidermis stripes. At doral closure, expression is absent from the amnioserosa but persists in the epidermis.
(Fernández et al., 2007)
Strong pMad staining is observed in the
(Shimmi et al., 2005)
precursor cells of the longitudinal veins around 19-22 hr after pupation. From 17-22 hr APF, pMAD accumulates in a broad patch of cells in the region of the presumptive posterior crossvein. During subsequent stages (13-28 hr APF), the pMAD staining becomes refined into a narrow stripe of cells that form the posterior crossvein.
(Shimmi et al., 2005)
Phosphorylated Mad protein is distributed in a gradient, with the highest levels near the anterior/posterior compartment boundary.
(Bornemann et al., 2004)
staining was for for pMad
(Gilboa and Lehmann, 2004)
In male adults, Mad protein is expressed in germline stem cells.
(Kawase et al., 2004)
In female adults, Mad protein is expressed at a high level in germline stem cells, and at a low level in cystoblasts.
(Song et al., 2004)
Phosphorylated Mad protein is localized to a subset of motor neurons in late stage embryos.
(Rawson et al., 2003)
Phosphorylated Mad (pMad) is first detected at future sites of the crossveins at 19-21 hr APF and in portions of the longitudinal veins near the crossvein attachment sites and along wing vein L2. By 24-26 hr APF staining is seen all along the longitudinal veins but is stronger near the crossveins. At 36 hr APF, staining is seen throughout the wing but is stronger in the crossveins. Strong staining is also observed in the nuclei and axons of the PNS of the wing at 18-36 hr APF.
(Conley et al., 2000)
Mad protein is ubiquitously distributed in embryos.
(Newfeld et al., 1996)
Marker for
 
crossvein
(Conley et al., 2000)
female germline stem cell
(Song et al., 2004)
germline stem cell
(Gilboa and Lehmann, 2004)
male germline stem cell
(Kawase et al., 2004)
Subcellular Localization
CV Term
Evidence
References
cytoplasm
inferred from direct assay
(Chen et al., 2006)
cytosol
inferred from direct assay
(Li and Li, 2006, Das et al., 1998, Wisotzkey et al., 1998, Newfeld et al., 1996)
heteromeric SMAD protein complex
inferred from physical interaction with FLYBASE:Med; FB:FBgn0011655
(Kamiya et al., 2008, Wisotzkey et al., 1998)
inferred from direct assay
(Inoue et al., 1998)
nucleus
inferred from direct assay
(Friedman et al., 2013, Chen et al., 2006, Li and Li, 2006, Das et al., 1998, Inoue et al., 1998, Wisotzkey et al., 1998, Kim et al., 1997)
inferred from direct assay
(assigned by UniProt )
(Colozza et al., 2011)
RNA polymerase II transcription repressor complex
inferred from physical interaction with FLYBASE:shn; FB:FBgn0003396, FLYBASE:Med; FB:FBgn0011655
(Yao et al., 2006, Muller et al., 2003)
transcription factor complex
inferred from physical interaction with FLYBASE:Med; FB:FBgn0011655
(Weiss et al., 2010)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Mad 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
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 4-6
  • Stages(s) 9-10
  • Stages(s) 11-12
FlyBase Wiki
Image Based Resources
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 53 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 44 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Mad
Transgenic constructs containing regulatory region of Mad
Deletions and Duplications ( 20 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
cell lethal | recessive | somatic clone
lethal (with Mad2)
lethal (with MadB1)
lethal (with Madk00223)
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 | recessive
lethal - all die before end of larval stage | recessive
lethal - all die before end of pupal stage, with Scer\GAL4da.G32
lethal - all die before end of pupal stage, with Scer\GAL4pnr-MD237
lethal - all die before end of pupal stage | recessive
lethal - all die before end of pupal stage | recessive | heat sensitive
lethal - all die during P-stage, with Scer\GAL4Mef2.PR
partially lethal - majority live, with Scer\GAL4Dll-md23
partially lethal - majority live, with Scer\GAL4how-24B
partially lethal - majority live, with Scer\GAL4T80
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4tin.CΔ4
viable (with Mad2L-112-1)
viable (with Mad2L-136-1)
viable (with Mad2L-163-23)
viable (with Mad6)
Sterility
Allele
female sterile (with Mad2L-112-1)
female sterile (with Mad2L-136-1)
female sterile (with Mad2L-163-23)
female sterile (with Mad6)
fertile
sterile | recessive | heat sensitive
Other Phenotypes
Allele
decreased cell number | adult stage | conditional, with Scer\GAL4VP16.nos.UTR
decreased cell number | adult stage | somatic clone
decreased cell number | somatic clone
decreased cell number | third instar larval stage, with Scer\GAL4repo.PU
decreased occurrence of cell division | larval stage | somatic clone
decreased occurrence of cell division | oogenesis | oogenesis
hyperplasia, with Scer\GAL4GMR24H06
hyperplasia | heat sensitive, with Scer\GAL4NP5130, Scer\GAL80ts.αTub84B
hyperplasia | somatic clone
increased cell death | third instar larval stage, with Scer\GAL4ap-md544
mitotic cell cycle defective | cell autonomous | somatic clone
neuroanatomy defective
neuroanatomy defective (with Df(2L)C28)
neuroanatomy defective (with Mad1)
neuroanatomy defective (with MadB65)
neuroanatomy defective | larval stage
neuroanatomy defective | third instar larval stage
neuroanatomy defective | third instar larval stage, with Scer\GAL4C57
neuroanatomy defective | third instar larval stage (with Mad1)
neuroanatomy defective | third instar larval stage (with Madk00237)
neurophysiology defective (with Mad1)
neurophysiology defective (with Mad12)
neurophysiology defective | third instar larval stage
short lived | RU486 conditional, with Scer\GAL4αTub84B.Switch.PK
size defective, with Scer\GAL4A9
size defective, with Scer\GAL4vg.MQ.Exel
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 | adult stage, with Scer\GAL4nub.PU
size defective | larval stage
small body | larval stage (with Mad1)
small body | larval stage (with MadB65)
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, with Scer\GAL4da.G32
some die during pupal stage, with Scer\GAL4pnr-MD237
some die during pupal stage | recessive
some die during pupal stage | recessive | heat sensitive
visible, with Scer\GAL4A9
visible, with Scer\GAL4ap-md544
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4Bx-MS1096-KE
visible, with Scer\GAL4ptc-559.1
visible, with Scer\GAL4sca-537.4
visible, with Scer\GAL4sd-SG29.1
visible, with Scer\GAL4vg.int2.1
visible, with Scer\GAL4vg.PM
visible | adult stage, with Scer\GAL4A9
visible | dominant
visible | embryonic stage, with Scer\GAL4VP16.mat.αTub67C
visible | heat sensitive (with Madapg)
visible | heat sensitive (with Madk00237)
visible | recessive
visible | recessive | somatic clone
visible | somatic clone
visible | somatic clone, with Scer\GAL4Act5C.PI
wild-type
Phenotype manifest in
Allele
abdominal segment
adult midgut progenitor cell | somatic clone
adult posterior midgut epithelium, with Scer\GAL4GMR24H06
adult posterior midgut epithelium | heat sensitive, with Scer\GAL4NP5130, Scer\GAL80ts.αTub84B
anterior wing margin | somatic clone, with Scer\GAL4Act5C.PI
apterous-expressing neuron of the lateral cluster of the thorax Ap4
bouton (with Df(2L)C28)
bouton | third instar larval stage, with Scer\GAL4C57
bouton | third instar larval stage, with Scer\GAL4futsch-C380
copper cell | somatic clone
crossvein, with Scer\GAL469B
crossvein | ectopic, with Scer\GAL4ptc-559.1
crystal cell
dense core granule | adult stage, with Scer\GAL4NP0261
denticle belt
denticle belt, with Scer\GAL4VP16.mat.αTub67C
denticle belt | ectopic, with Scer\GAL4VP16.mat.αTub67C
dorsal triple row | ectopic, with Scer\GAL4sd-SG29.1
dorsal triple row | supernumerary, with Scer\GAL4A9
dorsal triple row | supernumerary, with Scer\GAL4sd-SG29.1
egg
egg chamber
embryonic/first instar larval cuticle
embryonic/first instar larval cuticle | maternal effect
embryonic/larval central nervous system | larval stage
embryonic/larval dorsal branch
embryonic/larval fat body
embryonic/larval midgut | anterior
embryonic/larval neuromuscular junction | third instar larval stage
embryonic/larval neuromuscular junction | third instar larval stage, with Scer\GAL4C57
embryonic/larval neuromuscular junction | third instar larval stage, with Scer\GAL4futsch-C380
embryonic/larval neuromuscular junction | third instar larval stage (with Mad1)
embryonic/larval neuromuscular junction | third instar larval stage (with Madk00237)
embryonic/larval optic lobe
embryonic/larval somatic muscle (with Mad1)
embryonic/larval somatic muscle (with MadB65)
embryonic/larval tracheal system
embryonic/larval ventral anastomosis
eye
eye | somatic clone
eye-antennal disc
eye disc | cell autonomous | somatic clone
eye disc | posterior | somatic clone
eye disc | third instar larval stage, with Scer\GAL4repo.PU
fat body (with Mad7)
fat body (with Mad12)
female germline stem cell
female germline stem cell, with Scer\GAL4VP16.nos.UTR
female germline stem cell | adult stage | somatic clone
female germline stem cell | somatic clone
frons | ectopic | somatic clone
gastric caecum
germarium
germarium, with Scer\GAL4VP16.nos.UTR
histoblast | somatic clone
histoblast nest | somatic clone
imaginal disc
imaginal disc | maternal effect
interommatidial bristle | pupal stage P7 | somatic clone
intestinal stem cell | somatic clone
leg
leg | distal | heat sensitive (with Madapg)
leg | distal | heat sensitive (with Madk00237)
leg | ectopic, with Scer\GAL4ptc-559.1
leg | heat sensitive (with Madapg)
leg | heat sensitive (with Madk00237)
leg | 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
medial triple row | supernumerary, with Scer\GAL4A9
medial triple row | supernumerary, with Scer\GAL4sd-SG29.1
morphogenetic furrow
multivesicular body | adult stage, with Scer\GAL4dsx.KI
neuromuscular junction
neuromuscular junction, with Scer\GAL4Mhc.PW
neuromuscular junction (with Df(2L)C28)
neuromuscular junction (with Mad1)
neuromuscular junction (with MadB65)
neuromuscular junction | third instar larval stage (with Mad12)
neuromuscular junction | third instar larval stage (with Madk00237)
NMJ bouton
NMJ bouton | third instar larval stage
NMJ bouton | third instar larval stage (with Mad1)
NMJ bouton | third instar larval stage (with Madk00237)
nucleus | adult stage, with Scer\GAL4esg.PU
nucleus | somatic clone
ommatidium | somatic clone
ovariole
ovary, with Scer\GAL4VP16.nos.UTR
photoreceptor cell
photoreceptor cell | cell non-autonomous | somatic clone
photoreceptor cell | somatic clone
pleurum | somatic clone
posterior crossvein
secondary pigment cell | pupal stage P7 | somatic clone
secretory vesicle | adult stage, with Scer\GAL4dsx.KI
synapse
tarsal segment
tarsal segment | heat sensitive
tarsal segment | somatic clone
t-bar
tergite | somatic clone
terminal bouton (with Df(2L)C28)
terminal bouton | third instar larval stage (with Mad12)
terminal bouton | third instar larval stage (with Madk00237)
tertiary pigment cell | pupal stage P7 | somatic clone
testis | germline clone
tibia | heat sensitive
wing
wing, with Scer\GAL469B
wing, with Scer\GAL4A9
wing, with Scer\GAL4Bx-MS1096
wing, with Scer\GAL4nub.PU
wing, with Scer\GAL4nub-AC-62
wing, with Scer\GAL4ptc-559.1
wing, with Scer\GAL4sca-537.4
wing, with Scer\GAL4T80
wing, with Scer\GAL4vg.int2.1
wing, with Scer\GAL4vg.PM
wing | temperature conditional, with Scer\GAL4Bx-MS1096-KE
wing blade, with Scer\GAL4vg.MQ.Exel
wing cell
wing disc
wing disc | somatic clone
wing disc | third instar larval stage, with Scer\GAL4ap-md544
wing margin, with Scer\GAL469B
wing margin (with Mad11)
wing margin (with Mad12)
wing margin | somatic clone, with Scer\GAL4Act5C.PI
wing margin bristle, with Scer\GAL4A9
wing margin bristle | ectopic, with Scer\GAL4sca-537.4
wing margin bristle | ectopic, with Scer\GAL4sd-SG29.1
wing pouch, with Scer\GAL4sd-SG29.1
wing sensillum | ectopic, with Scer\GAL4A9
wing sensillum | ectopic, with Scer\GAL4sca-537.4
wing vein
wing vein, with Scer\GAL469B
wing vein, with Scer\GAL471B
wing vein, with Scer\GAL4A9
wing vein, with Scer\GAL4Bx-MS1096-KE
wing vein, with Scer\GAL4nub-AC-62
wing vein, with Scer\GAL4sca-537.4
wing vein, with Scer\GAL4vg.MQ.Exel
wing vein | ectopic, with Scer\GAL471B
wing vein | ectopic, with Scer\GAL4A9
wing vein | ectopic, with Scer\GAL4ptc-559.1
wing vein | ectopic, with Scer\GAL4sca-537.4
wing vein | ectopic, with Scer\GAL4vg.int2.1
wing vein L2 (with Mad11)
wing vein L2 (with Mad12)
wing vein L2 | cell autonomous | somatic clone
wing vein L3, with Scer\GAL4A9
wing vein L3 | cell autonomous | somatic clone
wing vein L4
wing vein L4, with Scer\GAL4A9
wing vein L4 (with Mad11)
wing vein L4 (with Mad12)
wing vein L5
wing vein L5, with Scer\GAL4A9
wing vein L5 (with Mad11)
wing vein L5 (with Mad12)
Orthologs
Downloads
Download All DIOPT Orthologs
Download All OrthoDB Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (9)
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\SMAD1
13 of 15
Yes
Yes
 
 
Hsap\SMAD5
10 of 15
No
Yes
Hsap\SMAD9
9 of 15
No
Yes
Hsap\SMAD2
2 of 15
No
No
 
Hsap\SMAD3
2 of 15
No
No
 
Hsap\SMAD4
1 of 15
No
No
 
Hsap\SMAD6
1 of 15
No
No
 
Hsap\SMAD7
1 of 15
No
No
 
Hsap\ZNF396
1 of 15
No
Yes
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\Smad1
13 of 15
Yes
Yes
Mmus\Smad5
9 of 15
No
Yes
Mmus\Smad9
9 of 15
No
Yes
Mmus\Smad2
2 of 15
No
No
Mmus\Smad3
2 of 15
No
No
Mmus\Smad4
1 of 15
No
No
Mmus\Smad6
1 of 15
No
No
Mmus\Smad7
1 of 15
No
No
Rattus norvegicus (Norway rat) (8)
Rnor\Smad1
11 of 13
Yes
Yes
Rnor\Smad5
9 of 13
No
Yes
Rnor\LOC103691556
5 of 13
No
Yes
Rnor\Smad9
5 of 13
No
Yes
Rnor\Smad2
2 of 13
No
No
Rnor\Smad3
2 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\smad1
10 of 12
Yes
Yes
Xtro\smad9
7 of 12
No
Yes
Xtro\smad2
1 of 12
No
No
Xtro\smad3
1 of 12
No
No
Xtro\smad4.1
1 of 12
No
No
Xtro\smad4.2
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
Danio rerio (Zebrafish) (13)
Drer\smad1
10 of 15
Yes
Yes
Drer\smad5
9 of 15
No
Yes
Drer\smad9
9 of 15
No
Yes
Drer\smad3a
2 of 15
No
No
Drer\smad3b
2 of 15
No
No
Drer\gb:cr929477
1 of 15
No
No
Drer\si:dkey-222b8.1
1 of 15
No
No
Drer\si:dkey-239n17.4
1 of 15
No
No
Drer\smad2
1 of 15
No
No
Drer\smad4a
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
Caenorhabditis elegans (Nematode, roundworm) (7)
Cele\sma-2
12 of 15
Yes
Yes
Cele\sma-3
8 of 15
No
Yes
Cele\daf-8
2 of 15
No
No
Cele\daf-14
2 of 15
No
No
Cele\daf-3
1 of 15
No
No
Cele\gei-4
1 of 15
No
Yes
Cele\tag-68
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (2)
Scer\RIM1
1 of 15
Yes
No
Scer\WHI3
1 of 15
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG09190715 )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Dmel\Mad
Drosophila suzukii
Spotted wing Drosophila
Dsuz\DS10_00000549
Drosophila simulans
Dsim\GD22774
Drosophila sechellia
Dsec\GM18164
Drosophila erecta
Dere\GG24457
Drosophila yakuba
Dyak\GE14935
Drosophila ananassae
Dana\GF20577
Drosophila pseudoobscura pseudoobscura
Dpse\GA28973
Drosophila persimilis
Dper\GL26531
Drosophila willistoni
Dwil\GK18268
Drosophila virilis
Dvir\GJ19553
Drosophila mojavensis
Dmoj\GI17981
Drosophila grimshawi
Dgri\GH10284
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091504EB )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Mdoa\17010300
Musca domestica
House fly
Mdoa\17026940
Glossina morsitans
Tsetse fly
Gmor\GMOY000441
Lucilia cuprina
Australian sheep blowfly
Lcup\FF38_14403
Mayetiola destructor
Hessian fly
Mdes\Mdes007752
Aedes aegypti
Yellow fever mosquito
Aaeg\AAEL005513
Anopheles darlingi
American malaria mosquito
Adar\ADAC002355
Anopheles gambiae
Malaria mosquito
Agam\Mad
Culex quinquefasciatus
Southern house mosquito
Cqui\CPIJ000420
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W04Z6 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bmor\BGIBMGA010481
Bombyx mori
Silkmoth
Bmor\BGIBMGA009650
Danaus plexippus
Monarch butterfly
Dple\DPOGS212475-PA
Danaus plexippus
Monarch butterfly
Dple\DPOGS202839-PA
Heliconius melpomene
Postman butterfly
Hmel\HMEL009327
Heliconius melpomene
Postman butterfly
Hmel\HMEL009295
Apis florea
Little honeybee
Aflo\2193906692
Apis florea
Little honeybee
Aflo\2417089716
Apis mellifera
Western honey bee
Amel\GB42213
Apis mellifera
Western honey bee
Amel\Ame.16140
Bombus impatiens
Common eastern bumble bee
Bimp\BIMP12830
Bombus impatiens
Common eastern bumble bee
Bimp\BIMP25611
Bombus terrestris
Buff-tailed bumblebee
Bter\15215919
Bombus terrestris
Buff-tailed bumblebee
Bter\15207820
Linepithema humile
Argentine ant
Lhum\LH12024
Linepithema humile
Argentine ant
Lhum\LH11668
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG007038
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG002089
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG012380
Dendroctonus ponderosae
Mountain pine beetle
Dpod\YQE_05291
Tribolium castaneum
Red flour beetle
Tcas\TC010162
Tribolium castaneum
Red flour beetle
Tcas\TC033446
Tribolium castaneum
Red flour beetle
Tcas\TC033447
Pediculus humanus
Human body louse
Phum\PHUM033930
Pediculus humanus
Human body louse
Phum\PHUM612490
Rhodnius prolixus
Kissing bug
Rpro\mad
Rhodnius prolixus
Kissing bug
Rpro\mad-like
Cimex lectularius
Bed bug
Clec\CLEC004107
Cimex lectularius
Bed bug
Clec\CLEC006591
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI010124
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI005545
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI007984
Zootermopsis nevadensis
Nevada dampwood termite
Znev\Znev_04615
Zootermopsis nevadensis
Nevada dampwood termite
Znev\Znev_07555
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X04JH )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Smar\SMAR008230
Strigamia maritima
European centipede
Smar\SMAR012602
Ixodes scapularis
Black-legged tick
Isca\ISCW010869
Stegodyphus mimosarum
African social velvet spider
Smim\KFM61359.1
Stegodyphus mimosarum
African social velvet spider
Smim\KFM64479.1
Tetranychus urticae
Two-spotted spider mite
Turt\tetur04g09040
Tetranychus urticae
Two-spotted spider mite
Turt\tetur11g03130
Daphnia pulex
Water flea
Dpux\DAPPUDRAFT_314805
Daphnia pulex
Water flea
Dpux\DAPPUDRAFT_300607
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G082C )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Smad2/3
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Smad1/5/8_1
Ciona intestinalis
Vase tunicate
Cint\ENSCING00000005942
Ciona intestinalis
Vase tunicate
Cint\ENSCING00000022986
Ciona intestinalis
Vase tunicate
Cint\ENSCING00000006188
Gallus gallus
Domestic chicken
Ggal\SMAD1
Gallus gallus
Domestic chicken
Ggal\ENSGALG00000014184
Gallus gallus
Domestic chicken
Ggal\SMAD5
Gallus gallus
Domestic chicken
Ggal\SMAD9
Gallus gallus
Domestic chicken
Ggal\SMAD3
Gallus gallus
Domestic chicken
Ggal\SMAD2
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (4)
Gene Symbol
Score
Source
Compara
eggNOG
Homologene
Inparanoid
Isobase
OrthoDB
OrthoFinder
orthoMCL
Panther
RoundUp
Alignment
Smox
5 of 10
Dad
3 of 10
Med
3 of 10
CG3719
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Allele
Disease
Evidence
References
Mad12
model of  juvenile polyposis syndrome
CEA
(Guo et al., 2013)
Potential Models Based on Orthology ( 1 )
Human Ortholog
Disease
Evidence
References
SMAD9; SMAD family member 9
model of  primary pulmonary hypertension
IEA
(FlyBase, 2019-)
Modifiers Based on Experimental Evidence ( 2 )
Allele
Disease
Interaction
References
Mad12
ameliorates  Angelman syndrome
modeled by Ube3a35
(Li et al., 2016)
ameliorates  muscular dystrophy
modeled by Scgδ840
(Goldstein et al., 2011)
DOES NOT ameliorate  cardiomyopathy
modeled by Scgδ840
(Goldstein et al., 2011)
MaddsRNA.IVS.13xlexAop
ameliorates  cancer
modeled by EgfrUAS.cBa, psqKK111691
(Herranz et al., 2014)
Comments on Models/Modifiers Based on Experimental Evidence ( 1 )
 
Mad12 suppresses muscle dysfunction but not the heart tube dysfunction of muscular dystrophy in Scgδ840 flies.
(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?
SMAD1; SMAD family member 1
13 of 15
601595
       
      SMAD5; SMAD family member 5
      10 of 15
      603110
          SMAD9; SMAD family member 9
          9 of 15
          603295
          Functional Complementation Data
          Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
          Dmel gene
          Ortholog showing functional complementation
          Supporting References
          Mad
          Hsap\SMAD1
          (Eivers et al., 2009)
          Interactions
          Summary of Physical Interactions
          esyN Network Diagram
          Show neighbor-neighbor interactions:
          Select Layout:
          Legend:
          Protein
          RNA
          Selected Interactor(s)
          Interactions Browser
          View in Interactions Browser
          Please see the Physical Interaction reports below for full details
          RNA-RNA
          Physical Interaction
          Assay
          References
          Mad - mir-124
          luminiscence technology
          (Sun et al., 2012)
          Mad - mir-ban
          luminiscence technology, necessary binding region
          (Robins et al., 2005)
          RNA-protein
          Physical Interaction
          Assay
          References
          Mad - bam
          anti tag coimmunoprecipitation, quantitative reverse transcription pcr
          (Malik et al., 2019)
          Mad - bgcn
          anti tag coimmunoprecipitation, quantitative reverse transcription pcr
          (Malik et al., 2019)
          Mad - brat
          anti tag coimmunoprecipitation, quantitative reverse transcription pcr
          (Malik et al., 2019)
          Mad - pum
          anti tag coimmunoprecipitation, quantitative reverse transcription pcr
          (Malik et al., 2019)
          protein-protein
          Physical Interaction
          Assay
          References
          Mad
          x-ray crystallography, molecular sieving, molecular weight estimation by staining, anti tag coimmunoprecipitation, anti tag western blot, two hybrid
          (Wang et al., 2009, Gao and Laughon, 2007, Inoue et al., 1998)
          Mad - Abd-B
          bimolecular fluorescence complementation, fluorescence microscopy
          (Baëza et al., 2015)
          Mad - Antp
          bimolecular fluorescence complementation, fluorescence microscopy
          (Baëza et al., 2015)
          Mad - Atf3
          two hybrid array
          (Shokri et al., 2019)
          Mad - Blos1
          two hybrid array
          (Shokri et al., 2019)
          Mad - CG14657
          anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based
          (Rhee et al., 2014)
          Mad - CG1738
          anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based
          (Rhee et al., 2014)
          Mad - CG4854
          two hybrid array
          (Shokri et al., 2019)
          Mad - CG7963
          two hybrid array
          (Shokri et al., 2019)
          Mad - CG8478
          two hybrid array
          (Shokri et al., 2019)
          Mad - Dd
          anti tag coimmunoprecipitation, western blot
          (Urrutia et al., 2016)
          Mad - Ets65A
          two hybrid array
          (Shokri et al., 2019)
          Mad - Hr78
          two hybrid array
          (Shokri et al., 2019)
          Mad - Irbp18
          two hybrid array
          (Shokri et al., 2019)
          Mad - Lis-1
          anti tag coimmunoprecipitation, anti tag western blot
          (Chen et al., 2010)
          Mad - MAN1
          pull down, autoradiography, two hybrid, anti bait coimmunoprecipitation, western blot
          (Wagner et al., 2010, Pinto et al., 2008)
          Mad - Med
          anti tag coimmunoprecipitation, anti tag western blot, pull down, tag visualisation, autoradiography
          (Cheng et al., 2013, Kazemian et al., 2013, Kamiya et al., 2008, Takaesu et al., 2006, Inoue et al., 1998, Wisotzkey et al., 1998)
          Mad - Pdp
          anti tag coimmunoprecipitation, anti tag western blot, pull down, western blot, enzymatic study
          (Chen et al., 2006)
          Mad - Rab5
          anti tag coimmunoprecipitation, anti tag western blot
          (Gui et al., 2016)
          Mad - Rel
          two hybrid array
          (Shokri et al., 2019)
          Mad - Scm
          two hybrid array
          (Shokri et al., 2019)
          Mad - Sec13
          anti tag coimmunoprecipitation, anti tag western blot, pull down, western blot
          (Chen and Xu, 2010)
          Mad - Tob
          anti tag coimmunoprecipitation, anti tag western blot
          (White-Grindley et al., 2014)
          Mad - Ubx
          bimolecular fluorescence complementation, fluorescence microscopy
          (Bischof et al., 2018, Baëza et al., 2015)
          Mad - abd-A
          bimolecular fluorescence complementation, fluorescence microscopy, anti tag coimmunoprecipitation, anti tag western blot
          (Bischof et al., 2018, Baëza et al., 2015)
          Mad - arm
          anti tag coimmunoprecipitation, anti tag western blot
          (Eivers et al., 2011)
          Mad - bin
          pull down, tag visualisation
          (Cheng et al., 2013)
          Mad - cnc
          two hybrid array
          (Shokri et al., 2019)
          Mad - eIF4A
          anti bait coimmunoprecipitation, western blot, anti tag coimmunoprecipitation, anti tag western blot, pull down
          (Li and Li, 2006)
          Mad - msk
          anti tag coimmunoprecipitation, western blot, anti tag western blot
          (Chen and Xu, 2010, Xu et al., 2007)
          Mad - nej
          pull down, autoradiography
          (Lilja et al., 2007)
          Mad - nmo
          anti tag coimmunoprecipitation, anti tag western blot, enzymatic study, autoradiography
          (Merino et al., 2009, Zeng et al., 2007)
          Mad - pan
          anti tag coimmunoprecipitation, anti tag western blot
          (Eivers et al., 2011, Zeng et al., 2008)
          Mad - pnr
          pull down, anti tag western blot, anti bait coimmunoprecipitation, western blot
          (Kim et al., 2017)
          Mad - ranshi
          two hybrid array
          (Shokri et al., 2019)
          Mad - rib
          pull down, tag visualisation
          (Cheng et al., 2013)
          Mad - scrib
          anti tag coimmunoprecipitation, anti tag western blot
          (Gui et al., 2016)
          Mad - seq
          two hybrid array
          (Shokri et al., 2019)
          Mad - shn
          pull down, autoradiography, anti tag coimmunoprecipitation, anti tag western blot, two hybrid
          (Dai et al., 2000)
          Mad - tkv
          anti tag coimmunoprecipitation, anti tag western blot, protein cross-linking with a bifunctional reagent, autoradiography
          (Gui et al., 2016, Kaneko et al., 2011, Inoue et al., 1998)
          Mad - vis
          anti tag coimmunoprecipitation, anti tag western blot
          (Hyman et al., 2003)
          Mad - yki
          anti tag coimmunoprecipitation, anti tag western blot, western blot
          (Oh and Irvine, 2011, Alarcón et al., 2009)
          Mad - zen
          pull down, autoradiography
          (Xu et al., 2005)
          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
          Mad
          suppressible
          Actβ
          (Gesualdi and Haerry, 2007)
          Mad
          suppressible
          anchor
          (Wang et al., 2018)
          Mad
          enhanceable
          babo
          (Gesualdi and Haerry, 2007)
          Mad
          suppressible
          Dad
          (Tsuneizumi et al., 1997)
          Mad
          suppressible
          eya
          (Curtiss and Mlodzik, 2000)
          Mad
          suppressible
          fuss
          (Fischer et al., 2012)
          Mad
          suppressible
          myo
          (Gesualdi and Haerry, 2007)
          Mad
          suppressible
          nmo
          (Zeng et al., 2007)
          Mad
          enhanceable
          put
          (Haerry, 2010)
          Mad
          enhanceable
          rl
          (Zeng et al., 2007)
          Mad
          suppressible
          Snoo
          (Barrio et al., 2007)
          Mad
          suppressible
          trio
          (Ball et al., 2010)
          Mad
          enhanceable
          trio
          (Ball et al., 2010)
          Mad
          suppressible
          wg
          (Eivers et al., 2011)
          Mad
          suppressible
          yki
          (Oh and Irvine, 2011)
          tkv|Mad
          enhanceable
          yki
          (Oh and Irvine, 2011)
          Starting gene(s)
          Interaction type
          Interacting gene(s)
          Reference
          anchor
          suppressible
          Mad
          (Wang et al., 2018)
          babo
          enhanceable
          Mad
          (Gesualdi and Haerry, 2007)
          babo
          suppressible
          Mad
          (Gesualdi and Haerry, 2007)
          brat
          suppressible
          Mad
          (Shi et al., 2013)
          brk
          suppressible
          Mad|tkv
          (Oh and Irvine, 2011)
          brk|tkv|yki
          enhanceable
          Mad
          (Oh and Irvine, 2011)
          cmpy
          suppressible
          Mad
          (James and Broihier, 2011)
          csw
          suppressible
          Mad
          (Oishi et al., 2006)
          CycE
          suppressible
          Mad|dpp
          (Horsfield et al., 1998)
          dally
          suppressible
          Mad
          (Kirkpatrick et al., 2006)
          Dd
          suppressible
          Mad
          (Liu et al., 2011)
          dpp
          enhanceable
          Mad
          (Takaesu et al., 2005, Raftery et al., 1995, Sekelsky et al., 1995)
          dpp
          suppressible
          Mad
          (Bangi and Wharton, 2006, Wiersdorff et al., 1996)
          Egfr|psq
          suppressible
          Mad
          (Herranz et al., 2014)
          ema
          suppressible
          Mad
          (Kim et al., 2010)
          gbb
          suppressible
          Mad
          (Bangi and Wharton, 2006)
          Neto
          enhanceable
          Mad
          (Sulkowski et al., 2014)
          Nlg4
          suppressible
          Mad
          (Zhang et al., 2017)
          Rac1
          suppressible
          Mad
          (Ball et al., 2010)
          S6KL
          suppressible
          Mad
          (Zhao et al., 2015)
          sax
          suppressible
          Mad
          (Waltzer and Bienz, 1999, Das et al., 1998)
          sca
          enhanceable
          Mad
          (Wiersdorff et al., 1996)
          Scgδ
          suppressible
          Mad
          (Goldstein et al., 2011)
          shg
          enhanceable
          Mad
          (Cordero et al., 2007)
          smo
          enhanceable
          Mad
          (Fu and Baker, 2003)
          Smox
          suppressible
          Mad
          (Hevia et al., 2017)
          Snoo
          suppressible
          Mad
          (Barrio et al., 2007, Takaesu et al., 2006)
          Snoo
          enhanceable
          Mad
          (Takaesu et al., 2006)
          sog
          enhanceable
          Mad
          (Decotto and Ferguson, 2001)
          spin
          suppressible
          Mad
          (Yuva-Aydemir et al., 2011)
          tkv
          suppressible
          Mad
          (Das et al., 1998, Hoodless et al., 1996, Wiersdorff et al., 1996)
          tor
          suppressible
          Mad
          (Li and Li, 2003)
          Ube3a
          suppressible
          Mad
          (Li et al., 2016)
          upd1
          suppressible
          Mad
          (Bach et al., 2003)
          Vps35
          suppressible
          Mad
          (Korolchuk et al., 2007)
          vvl
          enhanceable
          Mad
          (Certel and Johnson, 1998)
          yki
          suppressible
          Mad
          (Oh and Irvine, 2011, Reddy and Irvine, 2011)
          yki
          enhanceable
          Mad
          (Oh and Irvine, 2011)
          yki
          enhanceable
          Mad|tkv
          (Oh and Irvine, 2011)
          External Data
          Subunit Structure (UniProtKB)
          Homotrimer (PubMed:19557331). Interacts with MAN1 (PubMed:20036230). Interacts with Sec13 and Nup93-1 (PubMed:20547758).
          (UniProt, P42003 )
          Linkouts
          BioGRID - A database of protein and genetic interactions.
          DroID - A comprehensive database of gene and protein interactions.
          InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
          MIST (genetic) - An integrated Molecular Interaction Database
          MIST (protein-protein) - An integrated Molecular Interaction Database
          Pathways
          Gene Group - Pathway Membership (FlyBase)
          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.)
          External Data
          Linkouts
          KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
          SignaLink - A signaling pathway resource with multi-layered regulatory networks.
          Genomic Location and Detailed Mapping Data
          Chromosome (arm)
          2L
          Recombination map
          2-8
          Cytogenetic map
          23D3-23D3
          Sequence location
          2L:3,146,056..3,159,643 [-]
          FlyBase Computed Cytological Location
          Cytogenetic map
          Evidence for location
          23D3-23D3
          Limits computationally determined from genome sequence between P{PZ}toc01361 and P{lacW}Madk00237
          Experimentally Determined Cytological Location
          Cytogenetic map
          Notes
          References
          23D3-23D4
          (determined by in situ hybridisation)
          (Spradling et al., 1999)
          23D4-23D6
          (determined by in situ hybridisation)
          (Wiersdorff et al., 1996)
          23D-23D
          (determined by in situ hybridisation)
          (Roch et al., 1998, Newfeld et al., 1994)
          23D3-23D4
          (BDGP Project Members, 1994-1999)
          The Mad gene is missing in Df(2L)S2590, although the distal break of the cytologically defined deficiency falls in the toc gene.
          (Laverty, 1998.12.3)
          Experimentally Determined Recombination Data
          Location
          2-8
          (FlyBase, 2016)
          2-7.4
          (Comeron et al., 2012)
          2-7.7
          Left of (cM)
          (Decotto and Ferguson, 2001)
          Right of (cM)
          (Decotto and Ferguson, 2001)
          Notes
          Stocks and Reagents
          Stocks (28)
          Genomic Clones (44)
          cDNA Clones (130)
           

          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
          Other Information
          Relationship to Other Genes
          Source for database identify of
          Source for database merge of
          Source for merge of: E(zen)2 Mad
          (Ferguson, 1998.8.7)
          Source for merge of: Mad l(2)k00237
          (Laverty, 1998.12.3)
          Source for merge of: Mad apg
          (Galindo et al., 2002)
          Additional comments
          Other Comments
          DNA-protein interactions: genome-wide binding profile assayed for Mad protein in stage 9-11 embryos; ArrayExpress accession number E-MTAB-1184.
          (Junion et al., 2012)
          Phosphorylation of Mad protein in the linker region by sgg regulates the development of sensory organs in the anterior-dorsal quadrant of the wing.
          (Quijano et al., 2011)
          DNA-protein interactions: genome-wide binding profile assayed for Mad protein in 2-3 hr embryos; see BDTNP1_TFBS_Mad collection report.
          (MacArthur et al., 2009)
          The Mad product negatively regulates dac expression in the embryonic head.
          (Anderson et al., 2006)
          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)
          Hsap\MADH1 and Hsap\MADH4, as well as Mad and Med, can stimulate dpp signalling in limb development.
          (Marquez et al., 2001)
          Overexpression of Mad or Hsap\MADH1 in the wing and leg causes a similar phenotype.
          (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. Mad function is required to block salivary gland formation in dorsal regions of PS2.
          (Henderson et al., 1999)
          Mad may recruit nej to effect the transcriptional activation of dpp responsive genes during development.
          (Waltzer and Bienz, 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)
          Mutants show no interaction with Df(2R)Pcl11B or Df(3L)66C-G28.
          (Nicholls and Gelbart, 1998)
          Mutants isolated in a screen of the second chromosome identifying genes affecting disc morphology.
          (Roch et al., 1998)
          Mad mediated nuclear translocation is essential for Med function.
          (Wisotzkey et al., 1998)
          Transcriptional activation of Ubx is subject to competition between dpp-activated Mad and another Smad whose function as a transcriptional repressor depends on high wg signalling.
          (Yu et al., 1998)
          The amino-terminal domain of the Mad protein contains a sequence-specific DNA-binding activity that becomes apparent when carboxy-terminal residues are removed. Mad protein binds to and is required for the activation of an enhancer within the vg gene in cells across the entire developing wing blade.
          (Kim et al., 1997)
          Mad protein functions downstream of the serine-threonine kinase activity of the tkv receptor.
          (Newfeld et al., 1997)
          Signalling by constitutively active tkv mutation is suppressed by heterozygosity for Mad mutations. These results indicate that Mad functions downstream of the tkv receptor.
          (Hoodless et al., 1996)
          Mad is required for any response of the visceral mesoderm or endoderm to dpp signals from the visceral mesoderm and is required specifically in cells responding to dpp. Mad can function in the signalling pathway of BMP-4 in Xenopus embryos, and is thus a highly conserved and essential element of the dpp signalling pathway.
          (Newfeld et al., 1996)
          Mad is required for dpp signalling during eye development. Clonal analysis demonstrated that this requirement is cell autonomous. Mad is an essential component of the signal transduction pathway downstream of the dpp receptors in responding cells. Mad-mediated dpp signaling is absolutely required for the initiation of the morphogenetic furrow in the eye, but has only a minor role in its subsequent propagation across the eye disc.
          (Wiersdorff et al., 1996)
          Dominant enhancer of zen.
          (Hudson et al., 1995)
          Mad has been identified independently in two screens, one for dominant enhancers of the dpp mutant phenotype and dosage sensitive interactions with dpp. Mad mutant phenotypes show patterning defects that resemble many dpp mutant phenotypes. Studies of ectopic expression, tissue-specific expression and in situ hybridizations are consistent with a role for Mad downstream of dpp in the signalling pathway.
          (Newfeld et al., 1995)
          Genetic characterisation indicates Mad encodes a product essential for dpp function. Molecular analysis demonstrates the Mad protein is a member of a novel protein family that is highly conserved throughout metazoans.
          (Sekelsky et al., 1995)
          A genetic enhancer of dpp.
          (Jackson and Hoffmann, 1994)
          Loss of function mutations of Mad are dominant maternal effect enhancers of dpp during early embryogenesis and dominant zygotic enhancers of dpp in imaginal discs.
          (Newfeld et al., 1994)
          Origin and Etymology
          Discoverer
          Etymology
          The gene is named "apang" after the mutant phenotype ("apang" stands for an individual with mutilated and underdeveloped legs in Hindi).
          (Shekaran and Sharma, 1983)
          Identification
          External Crossreferences and Linkouts ( 54 )
          Sequence Crossreferences
          NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
          GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
          GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
          RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
          UniProt/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
          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.
          KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
          modMine - A data warehouse for the modENCODE project
          SignaLink - A signaling pathway resource with multi-layered regulatory networks.
          Linkouts
          ApoDroso - Functional genomic database for photoreceptor development, survival and function
          BioGRID - A database of protein and genetic interactions.
          DPiM - Drosophila Protein interaction map
          DroID - A comprehensive database of gene and protein interactions.
          DRSC - Results frm RNAi screens
          FLIGHT - Cell culture data for RNAi and other high-throughput technologies
          FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
          FlyMine - An integrated database for Drosophila genomics
          Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
          InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
          KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
          MIST (genetic) - An integrated Molecular Interaction Database
          MIST (protein-protein) - An integrated Molecular Interaction Database
          Synonyms and Secondary IDs (44)
          Reported As
          Symbol Synonym
          2/23
          (Roch et al., 1998)
          CG12399
          (Hevia et al., 2017, Neuert et al., 2017, Hosono et al., 2015, McEachern et al., 2014, Ni et al., 2011.7.19, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Ni et al., 2008, Brandt, 2006)
          E(zen)2
          En(vvl)
          MAD
          (Fan et al., 2018, Choudhury et al., 2017, Gleason et al., 2017, Liu and Jin, 2017, Grill et al., 2016, Blatti et al., 2015, Meinhardt, 2015, West et al., 2015, Halstead et al., 2014, Maimon et al., 2014, Minakhina et al., 2014, Restrepo et al., 2014, Slattery et al., 2014, Cheng et al., 2013, Humphreys et al., 2013, Li et al., 2013, Tremmel et al., 2013, Harvey and Hariharan, 2012, Mulligan et al., 2012, Niepielko et al., 2012, Chung et al., 2011, Goldstein et al., 2011, Niepielko et al., 2011, Szuperák et al., 2011, Chen and Xu, 2010, Haerry, 2010, Kang and Bier, 2010, Khuong et al., 2010, Mathur et al., 2010, Iovino et al., 2009, MacArthur et al., 2009, Nagaraj and Banerjee, 2009, Bickel et al., 2008, Ratnaparkhi et al., 2008, Rogulja et al., 2008, Yakoby et al., 2008, Fuller and Spradling, 2007, Gesualdi and Haerry, 2007, Potthoff and Olson, 2007, Shravage et al., 2007, Chen et al., 2006, Collins et al., 2006, Rusten et al., 2006, Cheesman et al., 2004, Kockel et al., 2001, Bassett et al., 1997)
          Mad
          (Chen, 2019, Kim et al., 2019, La Marca et al., 2019, Nakato et al., 2019, Shokri et al., 2019, Vuilleumier et al., 2019, Wang et al., 2019, Ahmed-de-Prado and Baonza, 2018, Ahmed-de-Prado et al., 2018, Ameku et al., 2018, Anreiter and Sokolowski, 2018, Bischof et al., 2018, Chandran et al., 2018, Levis, 2018.8.30, Liao et al., 2018, Newcomb et al., 2018, O'Hanlon et al., 2018, Powers and Srivastava, 2018, Prange et al., 2018, Roesley et al., 2018, Tseng et al., 2018, Wang et al., 2018, Aggarwal et al., 2017, Ahmad, 2017, Al Khatib et al., 2017, Banerjee et al., 2017, Follansbee et al., 2017, Golovnin et al., 2017, Houtz et al., 2017, Jeibmann et al., 2017, Jordán-Álvarez et al., 2017, Kim et al., 2017, Koenecke et al., 2017, Monsivais et al., 2017, Neuert et al., 2017, Requena et al., 2017, Richard et al., 2017, Romero-Pozuelo et al., 2017, Song et al., 2017, Tan et al., 2017, Transgenic RNAi Project members, 2017-, Zhou et al., 2017, Aggarwal et al., 2016, Bonfini et al., 2016, Djabrayan and Casanova, 2016, Fallahi et al., 2016, Frasch, 2016, Garaulet et al., 2016, Gui et al., 2016, Kwan et al., 2016, Li et al., 2016, Matsuda et al., 2016, Mbodj et al., 2016, Mottier-Pavie et al., 2016, Quijano et al., 2016, Sarov et al., 2016, Sharifkhodaei et al., 2016, Tauscher et al., 2016, Tsai et al., 2016, Urrutia et al., 2016, Akiyama and Gibson, 2015, Amoyel and Bach, 2015, Baëza et al., 2015, Ball et al., 2015, Bier and De Robertis, 2015, Bivik et al., 2015, Dolezal et al., 2015, Harmansa et al., 2015, Irvine and Harvey, 2015, Kakugawa et al., 2015, Kao et al., 2015, Kim et al., 2015, Liu et al., 2015, Newton et al., 2015, Sawala et al., 2015, Schertel et al., 2015, Schleede and Blair, 2015, Singh, 2015, Svendsen et al., 2015, Van Bortle et al., 2015, Yu et al., 2015, Zhou et al., 2015, Aleman et al., 2014, Alic et al., 2014, Archbold et al., 2014, Córdoba and Estella, 2014, Deshpande et al., 2014, Driver and Ohlstein, 2014, Esteves et al., 2014, Frank, 2014, Hamada-Kawaguchi et al., 2014, Herranz et al., 2014, Hodar et al., 2014, Jiang and Singh, 2014, Karandikar et al., 2014, Künnapuu et al., 2014, Navarro et al., 2014, Oh et al., 2014, Pichaud, 2014, Rhee et al., 2014, Rogers et al., 2014, Roy et al., 2014, Sulkowski et al., 2014, Tseng et al., 2014, White-Grindley et al., 2014, Wong et al., 2014, Zhang et al., 2014, Zhang et al., 2014, Bai et al., 2013, Bausek, 2013, Beckwith et al., 2013, Castellanos et al., 2013, Chang et al., 2013, Chauhan et al., 2013, Curtis et al., 2013, Doumpas et al., 2013, Enderle and McNeill, 2013, Guo et al., 2013, Ishitani and Ishitani, 2013, Jin et al., 2013, Jukam et al., 2013, Li and Gilmour, 2013, Li et al., 2013, Li et al., 2013, Marinho et al., 2013, Mbodj et al., 2013, Morozov and Ioshikhes, 2013, Nahm et al., 2013, Peterson and O'Connor, 2013, Saunders et al., 2013, Webber et al., 2013, Xin et al., 2013, Zhang et al., 2013, Aboukhalil and Bulyk, 2012, Agelopoulos et al., 2012, Akiyama et al., 2012, Beck et al., 2012, Boulanger et al., 2012, Cash and Andrews, 2012, Chen et al., 2012, Dahal et al., 2012, Dani et al., 2012, Fischer et al., 2012, Fuentes-Medel et al., 2012, Gomez et al., 2012, Kagey et al., 2012, Karim et al., 2012, Kim and Marqués, 2012, Le and Wharton, 2012, Lim et al., 2012, Liu et al., 2012, Pennetier et al., 2012, Peterson et al., 2012, Raftery and Umulis, 2012, Smith et al., 2012, Spokony and White, 2012.5.22, Stinchfield et al., 2012, Sun et al., 2012, Takaesu et al., 2012, Xia et al., 2012, Zhu et al., 2012, Abruzzi et al., 2011, Bayat et al., 2011, Ben-Zvi et al., 2011, Bhattacharya and Baker, 2011, Clark et al., 2011, de Cuevas and Matunis, 2011, Dejima et al., 2011, Dutko and Mullins, 2011, Eivers et al., 2011, Eliazer and Buszczak, 2011, Gancz et al., 2011, Giorgianni and Mann, 2011, Harris and Ashe, 2011, Issigonis and Matunis, 2011, König et al., 2011, Layalle et al., 2011, Li et al., 2011, Liu et al., 2011, Mikhaylova and Nurminsky, 2011, Miller et al., 2011, O'Keefe et al., 2011, Ogiso et al., 2011, Oh and Irvine, 2011, Peluso et al., 2011, Qian et al., 2011, Quijano et al., 2011, Rodal et al., 2011, Rodriguez, 2011, Veverytsa and Allan, 2011, Yang and Su, 2011, Yuva-Aydemir et al., 2011, Zhao et al., 2011, Ables and Drummond-Barbosa, 2010, Ballard et al., 2010, Ball et al., 2010, Chen et al., 2010, Ellis et al., 2010, Firth et al., 2010, Ishikawa et al., 2010, Li et al., 2010, Liu et al., 2010, Losada-Pérez et al., 2010, Mathur et al., 2010, Nahm et al., 2010, Neely et al., 2010, Padash-Barmchi et al., 2010, Portela et al., 2010, Quijano et al., 2010, Salzer et al., 2010, Sander et al., 2010, Sato et al., 2010, Su et al., 2010, Sun et al., 2010, Wagner et al., 2010, Wang and Ward, 2010, Weiss et al., 2010, Yagi et al., 2010, Alarcón et al., 2009, Eade and Allan, 2009, Eivers et al., 2009, Evans et al., 2009, Firth and Baker, 2009, Glavic et al., 2009, Guo and Wang, 2009, Guruharsha et al., 2009, Jensen et al., 2009, Künnapuu et al., 2009, Lembong et al., 2009, Liu et al., 2009, MacArthur et al., 2009, McKay et al., 2009, Merino et al., 2009, Pentek et al., 2009, Rhiner et al., 2009, Sellin et al., 2009, Smith-Bolton et al., 2009, Twombly et al., 2009, Umemori et al., 2009, Wang et al., 2009, Yu et al., 2009, Baena-Lopez et al., 2008, Bollenbach et al., 2008, Boylan et al., 2008, Brás-Pereira and Casares, 2008, Campbell and Moser, 2008, Casas-Tinto et al., 2008, Casas-Tinto et al., 2008, Chang et al., 2008, Christoforou et al., 2008, Erives et al., 2008, Estella and Mann, 2008, Estella et al., 2008, Fromental-Ramain et al., 2008, Fuentealba et al., 2008, Hao et al., 2008, Kamiya et al., 2008, Lembong et al., 2008, Lim et al., 2008, López-Onieva et al., 2008, Miguel-Aliaga et al., 2008, Miles et al., 2008, Newfeld et al., 2008, Ng, 2008, Pinto et al., 2008, Schwank et al., 2008, Serpe et al., 2008, Shcherbata et al., 2008, Shen et al., 2008, Takaesu et al., 2008, Tanaka-Matakatsu and Du, 2008, Tien et al., 2008, Tran and Doe, 2008, Ueyama et al., 2008, Vogler and Urban, 2008, Wang et al., 2008, Wang et al., 2008, Warrior et al., 2008, Yakoby et al., 2008, Yao et al., 2008, Yao et al., 2008, Zahedi et al., 2008, Zeng et al., 2008, Zhao et al., 2008, Aerts et al., 2007, Ambrus et al., 2007, Ballard and Wharton, 2007, Barrio et al., 2007, Baumgardt et al., 2007, Beltran et al., 2007, Christensen and Cook, 2007.3.22, Cordero et al., 2007, Crickmore and Mann, 2007, Escudero and Freeman, 2007, Fernández et al., 2007, Gao and Laughon, 2007, Gerlitz et al., 2007, Gonzales-Gaitan, 2007, Herrero et al., 2007, Hufnagel et al., 2007, Johnson et al., 2007, Kim et al., 2007, Korolchuk et al., 2007, Langille and Clark, 2007, Lecuit and Le Goff, 2007, Letizia et al., 2007, Liang et al., 2007, Li et al., 2007, Lilja et al., 2007, Makhijani et al., 2007, Manjon et al., 2007, Newfeld et al., 2007, Pistillo and Desplan, 2007, Ramel et al., 2007, Shcherbata et al., 2007, Shcherbata et al., 2007, Shirinian et al., 2007, Tyler and Baker, 2007, Tyler et al., 2007, Umemori et al., 2007, Walsh and Carroll, 2007, Xu et al., 2007, Yakoby et al., 2007, Zeng et al., 2007, Akasaka et al., 2006, Anderson et al., 2006, Bangi and Wharton, 2006, Bangi and Wharton, 2006, Bernardi et al., 2006, Bokel et al., 2006, Bokel et al., 2006, Bolivar et al., 2006, Brandt, 2006, Chen and Schupbach, 2006, Christensen and Cook, 2006.8.29, Christensen and Cook, 2006.8.29, Christensen and Cook, 2006.8.29, Christensen and Cook, 2006.12.5, Crickmore and Mann, 2006, Crickmore and Mann, 2006, Davidson and Erwin, 2006, de Navas et al., 2006, de Velasco et al., 2006, Dudu, 2006, Dudu et al., 2006, Dworkin and Gibson, 2006, Gao and Laughon, 2006, Garces and Thor, 2006, Jones et al., 2006, Joshi et al., 2006, Kirkpatrick et al., 2006, Knoblich, 2006, Layden et al., 2006, Li and Li, 2006, Lin et al., 2006, Mizutani et al., 2006, Niki, 2006, Niki et al., 2006, Oishi et al., 2006, Philippakis et al., 2006, Sotillos and de Celis, 2006, Takaesu et al., 2006, Umulis et al., 2006, van der Plas et al., 2006, Vrailas and Moses, 2006, Yao et al., 2006, Firth and Baker, 2005, Gao et al., 2005, Horabin, 2005, Kirilly et al., 2005, Marques, 2005, Mizutani et al., 2005, Sage et al., 2005, Stathopoulos and Levine, 2005, Takaesu et al., 2005, Xi and Xie, 2005, Yamashita et al., 2005, Koh et al., 2004, Pyrowolakis et al., 2004, Stanyon et al., 2004, Grienenberger et al., 2003, Jekely and Rorth, 2003, Rawson et al., 2003, Stefancsik and Sarkar, 2003, Wisotzkey et al., 2003, Saller and Bienz, 2001, Kyoda et al., 2000)
          Mat
          (Koh et al., 2004)
          P-Mad
          (Sotillos and De Celis, 2005)
          P-mad
          (López-Onieva et al., 2007)
          PMad
          (Wang et al., 2007)
          Smad
          (Itasaki and Hoppler, 2010, Leatherman and Dinardo, 2010)
          Smad1
          (Chen and Xu, 2010)
          apg
          c28
          (Wisotzkey et al., 2003)
          dMAD
          (Sorrentino et al., 2012, Kang and Bier, 2010)
          dMad
          (Hong et al., 2016, Hyman et al., 2003)
          l(2)K00237
          (Grammont et al., 1997)
          l(2)k00237
          mad
          (Lu et al., 2019, Malik et al., 2019, Whittle and Extavour, 2019, Kang et al., 2018, Laugks et al., 2017, Wells et al., 2017, Zhang et al., 2017, Lee et al., 2016, Li et al., 2016, Moulton and Letsou, 2016, Testa and Dworkin, 2016, Ayyaz et al., 2015, Charbonnier et al., 2015, Dong et al., 2015, Luo et al., 2015, Zhao et al., 2015, Kang et al., 2014, Liu et al., 2014, Atkins et al., 2013, Gafner et al., 2013, Jukam et al., 2013, Jukam et al., 2013, Shi et al., 2013, Chen et al., 2012, Fischer et al., 2012, Peterson et al., 2012, Xia et al., 2012, Dworkin et al., 2011, Giorgianni and Mann, 2011, Kaneko et al., 2011, Nègre et al., 2011, Reddy and Irvine, 2011, Vlachos and Harden, 2011, Wang et al., 2011, Kim et al., 2010, Ninov et al., 2010, Terriente-Félix et al., 2010, The modENCODE Consortium, 2010, The modENCODE Consortium, 2010, Vuilleumier et al., 2010, Debat et al., 2009, Twombly et al., 2009, Umemori et al., 2009, Widmann and Dahmann, 2009, Jiang et al., 2008, Ng, 2008, O'Connor-Giles et al., 2008, Seppa et al., 2008, Ulvklo et al., 2008, Zeng et al., 2008, Corrigall et al., 2007, Goold and Davis, 2007, Terriente and de Celis, 2007, Tyler and Baker, 2007, Dworkin and Gibson., 2006, Kirilly et al., 2006, Serpe and O'Connor, 2006, Eaton and Davis, 2005, Gibson and Perrimon, 2005, Kirilly et al., 2005, Li et al., 2005, Pappu et al., 2005, Xie et al., 2005, Dunlop et al., 2004, Kalinovsky and Scheiffele, 2004, Luschnig et al., 2004, Luschnig et al., 2004, Kawase and Xie, 2003, Kirilly et al., 2003, Shivdasani and Ingham, 2003, Zhu and Xie, 2003, Bessa et al., 2002, McCabe et al., 2002, Deng and Lin, 2001, Dorfman and Shilo, 2001, Affolter, 2000, Jordan et al., 2000, Marty et al., 2000, Zhang et al., 1999, Certel and Johnson, 1998, Cho and Blitz, 1998, Henderson and Andrew, 1998, Horsfield et al., 1998, Lin, 1998, Xie and Spradling, 1998, Yu et al., 1998, Henderson and Andrew, 1997, Mehler et al., 1997, Dean, 1996, Haerry et al., 1996, Hill, 1996, Hogan, 1996)
          p-MAD
          (Barrio and Milán, 2017)
          p-Mad
          (Adachi-Yamada et al., 2005)
          pMAD
          (Shilts and Broadie, 2017, Smelkinson et al., 2017, Friedman et al., 2013, König and Shcherbata, 2013, Bonds et al., 2007, Ward et al., 2006, Belenkaya et al., 2004, De Graeve et al., 2004, Kato et al., 2004)
          pMad
          (Fried et al., 2016, Raja et al., 2016, Monfort and Furlong, 2015.1.15, Sawala et al., 2015, Erceg et al., 2014, Pargett et al., 2014, Chang et al., 2013, Vanlandingham et al., 2013, Junion et al., 2012, Lu et al., 2012, Sopory et al., 2010, Sun et al., 2010, Umulis et al., 2010, Baumgardt et al., 2009, Guo and Wang, 2009, Nahmad and Stathopoulos, 2009, Yan et al., 2009, Boylan et al., 2008, Hatton-Ellis et al., 2007, Vrailas and Moses, 2006, Hatfield et al., 2005, Wang and Ferguson, 2005)
          pSmad
          (Leatherman and Dinardo, 2010)
          Name Synonyms
          Mother against DPP
          (Chauhan et al., 2013)
          Mother against Dpp
          (Fromental-Ramain et al., 2008)
          Mother against decapentaplegic
          (Vanlandingham et al., 2013, Manjon et al., 2007)
          Mothers Against Decapentaplegic
          (Paul et al., 2013, Dani et al., 2012, Higashi-Kovtun et al., 2010, Khuong et al., 2010)
          Mothers Against Dpp
          (Eliazer and Buszczak, 2011, Wang and Ward, 2010, Rusten et al., 2006)
          Mothers against DPP
          (Lembong et al., 2009, Bickel et al., 2008)
          Mothers against Decapentaplegic
          (Xu et al., 2007, Burnett et al., 2006, Chen et al., 2006)
          Mothers against Dpp
          (Hamada-Kawaguchi et al., 2014, Bausek, 2013, Chauhan et al., 2013, Jukam et al., 2013, König and Shcherbata, 2013, Agelopoulos et al., 2012, Lim et al., 2012, Harris and Ashe, 2011, Weiss et al., 2010, Evans et al., 2009, Umemori et al., 2009, Pinto et al., 2008, Zhao et al., 2008, Letizia et al., 2007, Anderson et al., 2006, Dworkin and Gibson, 2006, Horabin, 2005)
          Mothers against decapentaplegi
          (West et al., 2015)
          Mothers against decapentaplegic
          (Dani et al., 2012, Chen et al., 2010, Vrailas and Moses, 2006)
          Mothers against dpp
          (Djabrayan and Casanova, 2016, Mottier-Pavie et al., 2016, Li et al., 2014, McEachern et al., 2014, Roy et al., 2014, Xin et al., 2013, Aboukhalil and Bulyk, 2012, Liu et al., 2011, Ogiso et al., 2011, Ball et al., 2010, Losada-Pérez et al., 2010, Salzer et al., 2010, Twombly et al., 2009, Boylan et al., 2008, Chang et al., 2008, Bonds et al., 2007, Gao and Laughon, 2007, Umemori et al., 2007, Bernardi et al., 2006, Philippakis et al., 2006, Marques, 2005, Yamashita et al., 2005, Rawson et al., 2003, Stefancsik and Sarkar, 2003)
          Mothers-against-decapentaplegic
          (Svendsen et al., 2015)
          apang
          mothers against DPP
          (Harvey et al., 2013)
          mothers against decapentaplegi
          (Koh et al., 2004)
          mothers against decapentaplegic
          (Mathur et al., 2010, Lim et al., 2008)
          mothers against dpp
          (Humphreys et al., 2013, Firth et al., 2010, Terriente-Félix et al., 2010, Corrigall et al., 2007, Arbouzova and Zeidler, 2006)
          phospho-Smad
          (Leatherman and Dinardo, 2010)
          phosphoSmad
          (Leatherman and Dinardo, 2010)
          phosphorylated Smad
          (Baumgardt et al., 2009)
          Secondary FlyBase IDs
          • FBgn0000105
          • FBgn0013945
          • FBgn0022270
          • FBgn0024356
          Datasets (3)
          Study focus (3)
          Experimental Role
          Project
          Project Type
          Title
          • bait_protein
          Furlong_ChIP-chip
          ChIP-chip identification of binding sites for transcription factors that regulate mesodermal development.
          • 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 (1,023)