FB2025_05 , released December 11, 2025
Gene: Dmel\Mad
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General Information
Symbol
Dmel\Mad
Species
D. melanogaster
Name
Mothers against dpp
Annotation Symbol
CG12399
Feature Type
FlyBase ID
FBgn0011648
Gene Model Status
Stock Availability
Gene Summary
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] (FlyBase Gene Snapshot)
Also Known As

pMad, Mothers Against Decapentaplegic, Mother against decapentaplegic, dMad, Smad

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
2-8
RefSeq locus
NT_033779 REGION:3146056..3159643
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (47 terms)
Molecular Function (11 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:Med; FB:FBgn0288966
inferred from physical interaction with FLYBASE:vis; FB:FBgn0033748
inferred from physical interaction with UniProtKB:Q9V853
inferred from physical interaction with UniProtKB:Q02748
inferred from direct assay
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
Biological Process (30 terms)
Terms Based on Experimental Evidence (24 terms)
CV Term
Evidence
References
inferred from genetic interaction with FLYBASE:dpp; FB:FBgn0000490
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from expression pattern
inferred from direct assay
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
inferred from mutant phenotype
involved_in trunk segmentation
inferred from mutant phenotype
inferred from high throughput mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (8 terms)
CV Term
Evidence
References
Cellular Component (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
is_active_in cytosol
inferred from direct assay
inferred from direct assay
inferred from physical interaction with FLYBASE:Med; FB:FBgn0288966
is_active_in nucleus
inferred from direct assay
inferred from physical interaction with FLYBASE:Med; FB:FBgn0288966
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
Protein Family (UniProt)
Belongs to the dwarfin/SMAD family. (P42003)
Summaries
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]
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 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
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry P42003)

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

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

Experimentally Determined Structures
Crossreferences
PDB - An information portal to biological macromolecular structures
Comments on Gene Model

Gene model reviewed during 5.45

Gene model reviewed during 5.46

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0077616
2691
455
FBtr0332372
3262
525
Additional Transcript Data and Comments
Reported size (kB)

2.6 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0077302
50.5
455
7.04
FBpp0304648
57.9
525
7.20
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)
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
PDB - An information portal to biological macromolecular structures
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\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
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

-0.16

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: maternally deposited

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
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.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
organism | dorsal

Comment: Expressed in a dorsal midline stripe

primordial germ cell

Comment: antibody against phosphorylated form

organism | dorsal

Comment: antibody against phosphorylated form

primordial germ cell

Comment: antibody against phosphorylated form

female germline stem cell

Comment: high expression level

cystoblast

Comment: low expression level

in situ
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

pMad is expressed in CCAP-EN neurons but not in the CCAP-IN neurons.

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.

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.

Phosphorylated Mad protein is expressed in the presynaptic domain of type I boutons at the neuromuscular junction.

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.

Mad is observed in the dorsal epidermis and the ventrolateral epidermis at embryonic stage 13.

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.

Phosphorylated Mad protein is detected in centripetal follicle cells.

In third instar larval eye discs, the phosphorylated form of Mad is observed in a low-level, uniform granular distribution in and anterior to the morphogenetic furrow. Expression rises to a high level in regular groups of cells on the posterior side for two columns. This is followed by very low-level expression further posterior. The highest levels appear in groups of cells, just posterior to the first and second column of single ato-positive ommatidial founder cells and posterior to dpp transcription as revealed by a dpp lacZ reporter.

Strong pMad staining is observed in the

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.

Phosphorylated Mad protein is distributed in a gradient, with the highest levels near the anterior/posterior compartment boundary.

staining was for for pMad

In male adults, Mad protein is expressed in germline stem cells.

In female adults, Mad protein is expressed at a high level in germline stem cells, and at a low level in cystoblasts.

Phosphorylated Mad protein is localized to a subset of motor neurons in late stage embryos.

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.

Mad protein is ubiquitously distributed in embryos.

Marker for
Subcellular Localization
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
is_active_in cytosol
inferred from direct assay
inferred from direct assay
inferred from physical interaction with FLYBASE:Med; FB:FBgn0288966
is_active_in nucleus
inferred from direct assay
inferred from physical interaction with FLYBASE:Med; FB:FBgn0288966
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Mad in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 4-6
  • Stages(s) 9-10
  • Stages(s) 11-12
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 61 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 51 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Mad
Transgenic constructs containing regulatory region of Mad
Aberrations (Deficiencies and Duplications) ( 20 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (8)
13 of 14
Yes
Yes
 
1  
13 of 14
Yes
Yes
12 of 14
No
Yes
2  
5 of 14
No
No
1  
5 of 14
No
No
1  
2 of 14
No
No
8  
2 of 14
No
No
2  
2 of 14
No
No
1  
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (8)
13 of 14
Yes
Yes
12 of 14
No
Yes
10 of 14
No
Yes
5 of 14
No
No
5 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
Mus musculus (laboratory mouse) (8)
13 of 14
Yes
Yes
12 of 14
No
Yes
11 of 14
No
Yes
5 of 14
No
No
5 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
Xenopus tropicalis (Western clawed frog) (9)
11 of 13
Yes
Yes
10 of 13
No
Yes
9 of 13
No
Yes
4 of 13
No
No
3 of 13
No
No
2 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Danio rerio (Zebrafish) (13)
12 of 14
Yes
Yes
12 of 14
Yes
Yes
11 of 14
No
Yes
5 of 14
No
No
5 of 14
No
No
5 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (8)
13 of 14
Yes
Yes
4 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (4)
12 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Mad. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (5)
9 of 13
4 of 13
4 of 13
1 of 13
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 2 )
Potential Models Based on Orthology ( 1 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 4 )
Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
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
Interactions
Summary of Physical Interactions
Interaction Browsers

Please see the Physical Interaction reports below for full details
RNA-RNA
Physical Interaction
Assay
References
RNA-protein
Physical Interaction
Assay
References
protein-protein
Physical Interaction
Assay
References
Summary of Genetic Interactions
Interaction Browsers

Please look at the allele data for full details of the genetic interactions
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
enhanceable
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
suppressible
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.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Signaling Pathways (FlyBase)
Metabolic Pathways
FlyBase
External Links
External Data
Linkouts
KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
SignaLink - A signaling pathway resource with multi-layered regulatory networks.
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
2L
Recombination map
2-8
Cytogenetic map
Sequence location
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)
23D-23D
(determined by in situ hybridisation)
23D4-23D6
(determined by in situ hybridisation)
The Mad gene is missing in Df(2L)S2590, although the distal break of the cytologically defined deficiency falls in the toc gene.
Experimentally Determined Recombination Data
Location
Notes
Stocks and Reagents
Stocks (54)
Genomic Clones (44)
cDNA Clones (128)
 

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

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

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

cDNA Clones, End Sequenced (ESTs)
RNAi and Array Information
Linkouts
DRSC - Results frm RNAi screens
Antibody Information
Cell Line Information
Publicly Available Cell Lines
 
    Other Stable Cell Lines
     
      Other Comments

      DNA-protein interactions: genome-wide binding profile assayed for Mad protein in stage 9-11 embryos; ArrayExpress accession number E-MTAB-1184.

      Phosphorylation of Mad protein in the linker region by sgg regulates the development of sensory organs in the anterior-dorsal quadrant of the wing.

      DNA-protein interactions: genome-wide binding profile assayed for Mad protein in 2-3 hr embryos; see BDTNP1_TFBS_Mad collection report.

      The Mad product negatively regulates dac expression in the embryonic head.

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

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

      Hsap\MADH1 and Hsap\MADH4, as well as Mad and Med, can stimulate dpp signalling in limb development.

      Overexpression of Mad or Hsap\MADH1 in the wing and leg causes a similar phenotype.

      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.

      Mad may recruit nej to effect the transcriptional activation of dpp responsive genes during development.

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

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

      Mutants isolated in a screen of the second chromosome identifying genes affecting disc morphology.

      Mad mediated nuclear translocation is essential for Med function.

      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.

      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.

      Mad protein functions downstream of the serine-threonine kinase activity of the tkv receptor.

      Signalling by constitutively active tkv mutation is suppressed by heterozygosity for Mad mutations. These results indicate that Mad functions downstream of the tkv receptor.

      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.

      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.

      Dominant enhancer of zen.

      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.

      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.

      A genetic enhancer of dpp.

      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.

      Relationship to Other Genes
      Source for database merge of

      Source for merge of: E(zen)2 Mad

      Source for merge of: Mad l(2)k00237

      Source for merge of: Mad apg

      Additional comments
      Nomenclature History
      Source for database identify of
      Nomenclature comments
      Etymology

      The gene is named "apang" after the mutant phenotype ("apang" stands for an individual with mutilated and underdeveloped legs in Hindi).

      Synonyms and Secondary IDs (46)
      Reported As
      Symbol Synonym
      E(zen)2
      En(vvl)
      Mad
      (Kagemann et al., 2025, Tesson and Vincent, 2025, Al Asafen et al., 2024, Clémot et al., 2024, Eslahi et al., 2024, Jang et al., 2024, Nair and Baker, 2024, Shweta et al., 2024, Wang et al., 2024, Bastin and Eleftherianos, 2023, Cabrita and Martinho, 2023, Gou et al., 2023, Hossain et al., 2023, Lassetter et al., 2023, Liu et al., 2023, Matsuda and Affolter, 2023, Qian et al., 2023, Xue et al., 2023, Zhang et al., 2023, Zhou et al., 2023, Akiyama et al., 2022, Beaver et al., 2022, Bressan and Araujo, 2022, Chen et al., 2022, Enomoto and Igaki, 2022, Frampton et al., 2022, Montanari et al., 2022, Moore et al., 2022, Perlegos et al., 2022, Vicidomini and Serpe, 2022, Bahuguna et al., 2021, Boumard and Bardin, 2021, Connacher and Goldstrohm, 2021, Fabian et al., 2021, Hoshino and Niwa, 2021, Kochendoerfer et al., 2021, Kramer et al., 2021, Lin et al., 2021, Lodge et al., 2021, Madamanchi et al., 2021, Matsuda et al., 2021, Paul et al., 2021, Ruiz-Losada et al., 2021, Sidisky et al., 2021, Tian and Smith-Bolton, 2021, Yeates and Frank, 2021, Canales Coutiño et al., 2020, Cho et al., 2020, Chou et al., 2020, Cui et al., 2020, Frank et al., 2020, Gou et al., 2020, Hinnant et al., 2020, Jasper, 2020, Khan et al., 2020, Ladyzhets et al., 2020, Lamiré et al., 2020, Li et al., 2020, Mahmud et al., 2020, Moulton et al., 2020, Ote and Yamamoto, 2020, Pan et al., 2020, Port et al., 2020, Rushton et al., 2020, Schloop et al., 2020, Texada et al., 2020, Tu et al., 2020, Upadhyay et al., 2020, Zhou and Boutros, 2020, Berke et al., 2019, Chen, 2019, Chen and Zou, 2019, Drummond-Barbosa, 2019, Gui et al., 2019, Hoover et al., 2019, Kamimura et al., 2019, Kim et al., 2019, La Marca et al., 2019, Li and Baker, 2019, Li et al., 2019, Nakato et al., 2019, Politano et al., 2019, Shcherbata, 2019, Shokri et al., 2019, Svendsen et al., 2019, Vuilleumier et al., 2019, Wang et al., 2019, Williams-Simon et al., 2019, Ahmed-de-Prado and Baonza, 2018, Ahmed-de-Prado et al., 2018, Akiyama et 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      apg
      l(2)k00237
      mad
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      Name Synonyms
      Mathers against dpp
      Mother against DPP
      Mother against Dpp
      Mother against decapentaplegic
      Mothers against Decapentaplegic
      Mothers against decapentaplegi
      Mothers-against-decapentaplegic
      apang
      mothers against DPP
      mothers against decapentaplegi
      mothers against decapentaplegic
      phosphorylated Smad
      Secondary FlyBase IDs
      • FBgn0000105
      • FBgn0013945
      • FBgn0022270
      • FBgn0024356
      Datasets (3)
      Study focus (3)
      Experimental Role
      Project
      Project Type
      Title
      • bait_protein
      ChIP-chip identification of binding sites for transcription factors that regulate mesodermal development.
      • bait_protein
      ChIP characterization of transcription factor genome binding, Berkeley Drosophila Transcription Factor Network Project.
      • bait_protein
      Genome-wide localization of transcription factors by ChIP-chip and ChIP-Seq.
      Study result (0)
      Result
      Result Type
      Title
      External Crossreferences and Linkouts ( 59 )
      Sequence Crossreferences
      NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
      GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
      GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
      RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
      UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
      UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
      UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
      Other crossreferences
      AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
      BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
      DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
      EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
      FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
      FlyMine - An integrated database for Drosophila genomics
      KEGG Genes - Molecular building blocks of life in the genomic space.
      MARRVEL_MODEL - MARRVEL (model organism gene)
      PDB - An information portal to biological macromolecular structures
      Linkouts
      BioGRID - A database of protein and genetic interactions.
      Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
      DroID - A comprehensive database of gene and protein interactions.
      DRSC - Results frm RNAi screens
      Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
      FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
      FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
      Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
      Flygut - An atlas of the Drosophila adult midgut
      FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
      iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
      Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
      KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
      MIST (genetic) - An integrated Molecular Interaction Database
      MIST (protein-protein) - An integrated Molecular Interaction Database
      Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
      SignaLink - A signaling pathway resource with multi-layered regulatory networks.
      References (1,194)