FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\Dad
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General Information
Symbol
Dmel\Dad
Species
D. melanogaster
Name
Daughters against dpp
Annotation Symbol
CG5201
Feature Type
protein_coding_gene
FlyBase ID
FBgn0020493
Gene Model Status
Current
Stock Availability
32 publicly available
Gene Summary
Daughters against dpp (Dad) encodes the inhibitory SMAD in the BMP/Dpp pathway. It is involved in growth regulation and developmental patterning. [Date last reviewed: 2018-10-04] (FlyBase Gene Snapshot)
All Summaries
Also Known As

1883, l(3)j1E4

Key Links
Genomic Location
Cytogenetic map
89E11-89E11
Sequence location
Recombination map
3-59
RefSeq locus
NT_033777 REGION:17054007..17070408
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (27 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables BMP receptor binding
inferred from physical interaction with FLYBASE:tkv; FB:FBgn0003716
(Kamiya et al., 2008)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables I-SMAD binding
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
enables transcription regulator inhibitor activity
inferred from biological aspect of ancestor with PANTHER:PTN001008964
(GO Reference Genome Project, 2011-)
Biological Process (20 terms)
Terms Based on Experimental Evidence (15 terms)
CV Term
Evidence
References
involved_in anterior/posterior pattern specification, imaginal disc
inferred from mutant phenotype
(McClure and Schubiger, 2005)
involved_in cell elongation involved in imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Widmann and Dahmann, 2009)
involved_in cell migration
inferred from mutant phenotype
(Ninov et al., 2010)
involved_in eye-antennal disc development
inferred from mutant phenotype
(Stultz et al., 2006)
involved_in germ-line stem cell population maintenance
inferred from mutant phenotype
(Shivdasani and Ingham, 2003)
involved_in histoblast morphogenesis
inferred from mutant phenotype
(Ninov et al., 2010)
involved_in imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Dworkin and Gibson, 2006)
involved_in intestinal stem cell homeostasis
inferred from expression pattern
(Guo et al., 2013)
involved_in Malpighian tubule morphogenesis
inferred from mutant phenotype
(Bunt et al., 2010)
involved_in morphogenesis of larval imaginal disc epithelium
inferred from mutant phenotype
(McClure and Schubiger, 2005)
involved_in negative regulation of BMP signaling pathway
inferred from mutant phenotype
(Zhang et al., 2025, Liu et al., 2023, Ogiso et al., 2011, Kamiya et al., 2008, Tsuneizumi et al., 1997)
inferred from genetic interaction with FLYBASE:tkv; FB:FBgn0003716
(Tsuneizumi et al., 1997)
inferred from genetic interaction with FLYBASE:Mad; FB:FBgn0011648
(Tsuneizumi et al., 1997)
involved_in negative regulation of muscle organ development
inferred from mutant phenotype
(Jaramillo et al., 2009)
involved_in negative regulation of SMAD protein signal transduction
inferred from direct assay
(Kamiya et al., 2008, Inoue et al., 1998)
acts_upstream_of negative regulation of synaptic assembly at neuromuscular junction
inferred from mutant phenotype
(Nahm et al., 2010)
involved_in negative regulation of synaptic assembly at neuromuscular junction
inferred from mutant phenotype
(Nahm et al., 2013)
involved_in retinal ganglion cell axon guidance
inferred from mutant phenotype
(Yoshida et al., 2005)
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
involved_in anatomical structure morphogenesis
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
involved_in cell differentiation
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
involved_in regulation of DNA-templated transcription
(InterPro Project Members, 2004-)
involved_in regulation of transcription by RNA polymerase II
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
involved_in SMAD protein signal transduction
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
Cellular Component (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in cytosol
inferred from direct assay
(Liu et al., 2023, Sharifkhodaei and Auld, 2021)
located_in nucleus
inferred from direct assay
(Liu et al., 2023, Sharifkhodaei and Auld, 2021)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
part_of heteromeric SMAD protein complex
inferred from biological aspect of ancestor with PANTHER:PTN000344419
(GO Reference Genome Project, 2011-)
part_of transcription regulator complex
inferred from electronic annotation with InterPro:IPR013019
(InterPro Project Members, 2004-)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
Daughters against dpp (Dad) encodes the inhibitory SMAD in the BMP/Dpp pathway. It is involved in growth regulation and developmental patterning. [Date last reviewed: 2018-10-04]
Pathway (FlyBase)
NEGATIVE REGULATORS OF BMP SIGNALING PATHWAY -
Negative regulators of Bone Morphogenetic Protein (BMP) signaling down-regulate the pathway, ultimately resulting in the decreased nuclear activity of the Mad/Med transcription factor complex.
Summary (Interactive Fly)

Smad family - inhibits Dpp signal transduction - Ubx dynamically regulates Dpp signaling by repressing Dad expression during copper cell regeneration in the adult Drosophila midgut

Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
2

Please see the JBrowse view of Dmel\Dad 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 A0A6H2EEY2)

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

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

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

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

Gene model reviewed during 5.49

Stop-codon suppression (UGA) postulated; FBrf0243886.

Gene model reviewed during 6.32

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Dad-RA
FBtr0083393
NM_057912
3767
568
Dad-RB
FBtr0083394
NM_169740
3550
568
Dad-RC
FBtr0100160
NM_001014629
3751
568
Dad-RD
FBtr0479820
NM_001382177
3751
665
Additional Transcript Data and Comments
Reported size (kB)

3.8 (longest cDNA)

(Tsuneizumi et al., 1997)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
Dad-PA
FBpp0082834
63.6
568
7.87
Q5U110
NP_477260
AAN13728
Dad-PB
FBpp0082835
63.6
568
7.87
Q5U110
NP_732196
AAF55383
Dad-PC
FBpp0099512
63.6
568
7.87
Q5U110
NP_001014629
AAX52959
Dad-PD
FBpp0428153
74.7
665
8.69
QJC18471
Polypeptides with Identical Sequences

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

568 aa isoforms: Dad-PA, Dad-PB, Dad-PC
Additional Polypeptide Data and Comments
Reported size (kDa)

568 (aa); 63.6 (kD predicted)

(Tsuneizumi et al., 1997)
Comments
External Data
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\Dad using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Testis-specificity index

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

-1.78

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 11
dorsal ectoderm
(Marty et al., 2000)
third instar larval stage
wing disc | restricted
(Tsuneizumi et al., 1997)
wandering third instar larval stage
anterior-posterior compartment boundary of the wing disc
(Tsuneizumi et al., 1997)
Additional Descriptive Data

Dad transcript is expressed at the A/P boundary of the wing disc, in a stripe overlapping but broader than that of dpp.

(Tsuneizumi et al., 1997)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytosol
inferred from direct assay
(Liu et al., 2023, Sharifkhodaei and Auld, 2021)
located_in nucleus
inferred from direct assay
(Liu et al., 2023, Sharifkhodaei and Auld, 2021)
Expression Deduced from Reporters
Reporter: P{Dad-EGFP.nuclear}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 13 -- 16
embryonic/larval anterior Malpighian tubule | restricted
(Weavers and Skaer, 2013)
Reporter: P{Dad-GFP.N}
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
air sac primordium

Comment: wing disc associated trachea

(Roy et al., 2014)
pupal stage P5 -- P6
histoblast nest | restricted

Comment: periphery, 18-19 hr APF

(Ninov et al., 2010)
Reporter: P{Dad-lacZ.N}
Stage
Tissue/Position (including subcellular localization)
Reference
pupal stage P5 -- P6
histoblast nest | restricted

Comment: periphery, 18-19 hr APF

(Ninov et al., 2010)
Reporter: P{lacW}Dadj1E4
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Tsuneizumi et al., 1997)
larval stage
larval ventral nerve cord | restricted
(Vuilleumier et al., 2019)
third instar larval stage
wing disc | restricted
(Tsuneizumi et al., 1997)
larval ventral nerve cord
(Vuilleumier et al., 2019)
adult stage
adult midgut
(Guo et al., 2013)
adult midgut region R3a
(Guo et al., 2013)
adult midgut region R3b
(Guo et al., 2013)
adult s-LNv neuron
(Beckwith et al., 2013)
DN1 neuron | subset
(Beckwith et al., 2013)
Reporter: P{lacZ}DadP1883
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Tsuneizumi et al., 1997)
third instar larval stage
wing disc | restricted
(Tsuneizumi et al., 1997)
eye disc | restricted
(Firth et al., 2010)
wandering third instar larval stage
anterior-posterior compartment boundary of the wing disc
(Tsuneizumi et al., 1997)
oogenesis
female germline stem cell
(Song et al., 2004)
cystoblast | subset
(Song et al., 2004)
oogenesis stage S8 -- S10
centripetal follicle cell
(Muzzopappa and Wappner, 2005)
oogenesis stage S10
border follicle cell
(Muzzopappa and Wappner, 2005)
spermatogenesis
male germline stem cell
(Kawase et al., 2004)
primary gonial cell
(Kawase et al., 2004)
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Dad in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 40 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 17 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Dad
Transgenic constructs containing regulatory region of Dad
Aberrations (Deficiencies and Duplications) ( 5 )
Inferred from experimentation ( 5 )
Gene not disrupted in
Df(3R)C4
(Spradling et al., 1999)
Df(3R)ED5780
(Ryder and Roote, 2004.11.10)
Gene disrupted in
Df(3)DadP1883Δ32
(Bieli et al., 2015)
Df(3R)Exel6176
(Ryder, 2004.4.26)
Gene partially disrupted in
T(2;3)apXa
(Bieli et al., 2015)
Inferred from location ( 6 )
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
cell lethal | somatic clone, with Scer\GAL4Act5C.PP
lethal, with Scer\GAL432B
lethal, with Scer\GAL469B
lethal, with Scer\GAL4ap-md544
lethal, with Scer\GAL4C-765
lethal, with Scer\GAL4Dll-md23
lethal, with Scer\GAL4dpp.blk1
lethal, with Scer\GAL4en-e16E
lethal, with Scer\GAL4how-24B
lethal, with Scer\GAL4pnr-MD237
lethal, with Scer\GAL4ptc-559.1
lethal, with Scer\GAL4T80
lethal | recessive
partially lethal - majority die
partially lethal - majority die | recessive
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4tin.CΔ4
Other Phenotypes
Allele
abnormal cell shape, with Scer\GAL4Ubx-lac1-GAL4
abnormal neuroanatomy, with Scer\GAL4RapGAP1-OK6
abnormal neuroanatomy | adult stage, with Scer\GAL4elav-C155
abnormal neuroanatomy | larval stage
abnormal neuroanatomy | larval stage (with Dad271-68)
abnormal neuroanatomy | larval stage (with Dadj1E4)
abnormal neuroanatomy | recessive
abnormal neuroanatomy | third instar larval stage
abnormal neurophysiology, with Scer\GAL4Mhc.PW
abnormal pain response | adult stage, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
abnormal size | adult stage, with Scer\GAL4dsx.KI
abnormal size | adult stage, with Scer\GAL4esg.PU
abnormal size | adult stage, with Scer\GAL4NP0261
abnormal size | larval stage, with Scer\GAL4A9
decreased cell growth rate | adult stage | virgin male, with Scer\GAL4esg.PU
decreased occurrence of cell division | adult stage | conditional, with Scer\GAL4NP0001
decreased occurrence of cell division | cell autonomous | somatic clone | third instar larval stage, with Scer\GAL4Act5C.Switch.PR
decreased rate of larval locomotory behavior
hypoplasia | somatic clone
increased cell death | somatic clone, with Scer\GAL4Act5C.PI
increased occurrence of cell division | cell non-autonomous | somatic clone | third instar larval stage, with Scer\GAL4Act5C.Switch.PR
short lived | heat sensitive, with Scer\GAL4tin.CΔ4
visible, with Scer\GAL4A9
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4dpp.3KK
visible, with Scer\GAL4dpp.s-hc
visible, with Scer\GAL4vg.PM
visible | adult stage | male
visible | dominant
Phenotype manifest in
Allele
adult brain, with Scer\GAL4elav-C155
adult midgut | conditional, with Scer\GAL4NP0001
amnioserosa, with Scer\GAL4esg.FC
anterior crossvein, with Scer\GAL471B
bouton | increased number
dense core granule | adult stage, with Scer\GAL4NP0261
dorsal appendage
dorsal appendage, with Scer\GAL4CY2
dorsal appendage, with Scer\GAL4GR1
dorsal appendage | ectopic
dorsal tracheal branch primordium, with Scer\GAL4btl.PS
egg chamber, with Scer\GAL4GR1
egg operculum, with Scer\GAL4CY2
embryonic/larval neuromuscular junction | third instar larval stage
embryonic anterior Malpighian tubule, with Scer\GAL4byn-Gal4
embryonic dorsal epidermis, with Scer\GAL4esg.FC
extracellular exosome, with Scer\GAL4dsx.KI
eye disc & peripodial epithelium, with Scer\GAL4c311
eye photoreceptor cell stalk, with Scer\GAL4bi-omb-Gal4
fat body, with Scer\GAL4Act5C.PI
female germline stem cell
female germline stem cell, with Scer\GAL4nanos.PG
female germline stem cell, with Scer\GAL4VP16.nanos.UTR
filamentous actin, with Scer\GAL4esg.FC
germline cell | female | larval stage, with Scer\GAL4VP16.nanos.UTR
germline cyst | ectopic, with Scer\GAL4VP16.nanos.UTR
histoblast | somatic clone, with Scer\GAL4FRT.Act5C
histoblast nest | somatic clone, with Scer\GAL4FRT.Act5C
intestinal stem cell | somatic clone
lamina, with Scer\GAL4bi-omb-Gal4
lamina anlage glial cell, with Scer\GAL4bi-omb-Gal4
lamina intrinsic neuron Lai, with Scer\GAL4bi-omb-Gal4
lamina tangential neuron, with Scer\GAL4bi-omb-Gal4
leg disc, with Scer\GAL4Dll-md23
lysosome | adult stage, with Scer\GAL4dsx.KI
lysosome | adult stage, with Scer\GAL4NP0261
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, with Scer\GAL4VP16.nanos.UTR
maxillary palp, with Scer\GAL4dpp.s-hc
mesothoracic tergotrochanter muscle cell, with Scer\GAL41151
multivesicular body, internal vesicle | increased number, with Scer\GAL4dsx.KI
multivesicular body | adult stage, with Scer\GAL4dsx.KI
multivesicular body | adult stage, with Scer\GAL4NP0261
muscle cell of A1-7 lateral longitudinal muscle 1 | third instar larval stage
neuromuscular junction
neuromuscular junction, with Scer\GAL4RapGAP1-OK6
NMJ bouton
NMJ bouton | increased number | third instar larval stage
NMJ bouton | larval stage | increased number
NMJ bouton | larval stage | increased number (with Dad271-68)
NMJ bouton | larval stage | increased number (with Dadj1E4)
NMJ bouton | third instar larval stage
nucleus | adult stage, with Scer\GAL4dsx.KI
nucleus | adult stage, with Scer\GAL4esg.PU
nucleus | adult stage, with Scer\GAL4NP0261
nurse cell, with Scer\GAL4GR1
peripodial epithelium, with Scer\GAL4Ubx-lac1-GAL4
spermatocyte, with Scer\GAL4VP16.nanos.UTR
spermatogonium, with Scer\GAL4VP16.nanos.UTR
stretch follicle cell, with Scer\GAL4GR1
testis, with Scer\GAL4VP16.nanos.UTR
thoracic adult muscle precursor cell, with Scer\GAL41151
tracheal tip cell | embryonic stage, with Scer\GAL4btl.PS
vibrissal bristle, with Scer\GAL4dpp.s-hc
wing
wing, with Scer\GAL4Bx-MS1096
wing, with Scer\GAL4Gug-AGiR
wing, with Scer\GAL4nub-AC-62
wing, with Scer\GAL4salm-459.2
wing, with Scer\GAL4vg.PM
wing | male
wing disc, with Scer\GAL4A9
wing disc, with Scer\GAL4Dll-md23
wing disc, with Scer\GAL4Gug-AGiR
wing disc, with Scer\GAL4nub-AC-62
wing disc, with Scer\GAL4Ubx-lac1-GAL4
wing disc | cell autonomous | third instar larval stage
wing disc | cell non-autonomous | medial | third instar larval stage
wing disc | somatic clone, with Scer\GAL4Act5C.PI
wing margin, with Scer\GAL4Gug-AGiR
wing pouch, with Scer\GAL4en-e16E
wing vein
wing vein, with Scer\GAL4dpp.3KK
wing vein, with Scer\GAL4salm-459.2
wing vein | ectopic
wing vein L2
wing vein L4, with Scer\GAL471B
wing vein L5, with Scer\GAL471B
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (8)
Hsap\SMAD6
9 of 14
Yes
Yes
2  
Hsap\SMAD7
8 of 14
No
Yes
1  
Hsap\SMAD4
3 of 14
No
No
8  
Hsap\SMAD1
2 of 14
No
No
1  
Hsap\SMAD2
2 of 14
No
No
1  
Hsap\SMAD3
2 of 14
No
No
1  
Hsap\SMAD5
2 of 14
No
No
2  
Hsap\SMAD9
2 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (8)
Rnor\Smad6
10 of 14
Yes
Yes
Rnor\Smad7
9 of 14
No
Yes
Rnor\Smad1
2 of 14
No
No
Rnor\Smad2
2 of 14
No
No
Rnor\Smad3
2 of 14
No
No
Rnor\Smad4
2 of 14
No
No
Rnor\Smad5
2 of 14
No
No
Rnor\Smad9
1 of 14
No
No
Mus musculus (laboratory mouse) (8)
Mmus\Smad6
9 of 14
Yes
Yes
Mmus\Smad7
9 of 14
Yes
Yes
Mmus\Smad4
3 of 14
No
No
Mmus\Smad1
2 of 14
No
No
Mmus\Smad2
2 of 14
No
No
Mmus\Smad3
2 of 14
No
No
Mmus\Smad5
2 of 14
No
No
Mmus\Smad9
2 of 14
No
No
Xenopus tropicalis (Western clawed frog) (10)
Xtro\smad6
6 of 13
Yes
Yes
Xtro\smad7
6 of 13
Yes
Yes
Xtro\aldh7a1
1 of 13
No
No
Xtro\rpl14
1 of 13
No
No
Xtro\smad1
1 of 13
No
No
Xtro\smad2
1 of 13
No
No
Xtro\smad6.2
1 of 13
No
Yes
Xtro\smad9
1 of 13
No
No
Xtro\tmem54
1 of 13
No
No
Xtro\tspan15
1 of 13
No
No
Danio rerio (Zebrafish) (13)
Drer\smad6a
9 of 14
Yes
Yes
Drer\smad6b
9 of 14
Yes
Yes
Drer\smad7
8 of 14
No
Yes
Drer\smad4a
3 of 14
No
No
Drer\smad2
2 of 14
No
No
Drer\smad3a
2 of 14
No
No
Drer\smad3b
2 of 14
No
No
Drer\smad5
2 of 14
No
No
Drer\smad10a
2 of 14
No
No
Drer\gb:cr929477
1 of 14
No
No
Drer\smad1
1 of 14
No
No
Drer\smad4b
1 of 14
No
No
Drer\smad9
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (8)
Cele\tag-68
5 of 14
Yes
Yes
Cele\sma-3
3 of 14
No
No
Cele\daf-8
2 of 14
No
No
Cele\sma-2
2 of 14
No
No
Cele\C34E11.2
1 of 14
No
No
Cele\daf-3
1 of 14
No
No
Cele\daf-14
1 of 14
No
No
Cele\sma-4
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (4)
Agam\AgaP_AGAP002902
2 of 12
Yes
No
Agam\AgaP_AGAP008551
2 of 12
Yes
No
Agam\AgaP_AGAP009777
2 of 12
Yes
No
Agam\AgaP_AGAP007416
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Dad. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (5)
Mad
4 of 13
Med
4 of 13
Smox
4 of 13
exp
1 of 13
reb
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 3 )
    Human Ortholog
    Disease
    Evidence
    References
    SMAD6; SMAD family member 6
    model of  craniosynostosis 7
    IEA
    (FlyBase, 2019-)
    model of  aortic valve disease 2
    IEA
    (FlyBase, 2019-)
    model of  radioulnar synostosis
    IEA
    (FlyBase, 2019-)
    Modifiers Based on Experimental Evidence ( 3 )
    Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    SMAD6; SMAD family member 6
    9 of 14
    602931
     
    SMAD7; SMAD family member 7
    8 of 14
    602932
       
      SMAD4; SMAD family member 4
      3 of 14
      SMAD FAMILY MEMBER 4; SMAD4
       
      Functional Complementation Data
      Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
      Interactions
      Summary of Physical Interactions
      Summary of Genetic Interactions
      Interaction Browsers
      View in FlyBase Interactions Browser   View in MIST tool (external link)
      Please look at the allele data for full details of the genetic interactions
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      Dad
      suppressible
      arm
      (Ribeiro et al., 2004)
      Dad
      suppressible
      Art1
      (Xu et al., 2013)
      Dad
      suppressible
      Cln7
      (Connolly et al., 2019)
      Dad
      suppressible
      Dap160
      (O'Connor-Giles et al., 2008)
      Dad
      enhanceable
      LamC
      (Mohar et al., 2025)
      Dad
      suppressible
      Lpin
      (Liu et al., 2011)
      Dad
      suppressible
      Mbs
      (Widmann and Dahmann, 2009)
      Dad
      suppressible
      Nlg4
      (Zhang et al., 2017)
      Dad
      suppressible
      nwk
      (O'Connor-Giles et al., 2008)
      Dad
      suppressible
      Rho1
      (Widmann and Dahmann, 2009)
      Dad
      suppressible
      sqh
      (Widmann and Dahmann, 2009)
      Dad
      suppressible
      tkv
      (Ogiso et al., 2011)
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      Acsl
      enhanceable
      Dad
      (Liu et al., 2014)
      brk
      suppressible
      Dad
      (Martin et al., 2004)
      Cdk4|CycD
      suppressible
      Dad
      (Sekar et al., 2023)
      Cln7
      suppressible
      Dad
      (Connolly et al., 2019)
      Ctdnep1
      enhanceable
      Dad
      (Liu et al., 2011)
      Ctdnep1
      suppressible
      Dad
      (Liu et al., 2011)
      cup
      enhanceable
      Dad
      (Menon et al., 2015)
      CycE
      enhanceable
      Dad
      (Horsfield et al., 1998)
      dally
      suppressible
      Dad
      (Guo and Wang, 2009)
      Delta
      suppressible
      Dad
      (Ward et al., 2006)
      Dg
      enhanceable
      Dad
      (Kucherenko et al., 2008)
      Dys
      suppressible
      Dad
      (Kucherenko et al., 2008)
      fu
      suppressible
      Dad
      (Xia et al., 2010)
      gbb
      suppressible
      Dad
      (Kawase et al., 2004)
      Gli
      suppressible
      Dad
      (Sharifkhodaei et al., 2016)
      Gli
      enhanceable
      Dad
      (Sharifkhodaei et al., 2016)
      H
      enhanceable
      Dad
      (Johannes and Preiss, 2002)
      L
      enhanceable
      Dad
      (Singh et al., 2005)
      LamC
      suppressible
      Dad
      (Mohar et al., 2025)
      LamC
      enhanceable
      Dad
      (Mohar et al., 2025)
      lili
      suppressible
      Dad
      (Dolezal et al., 2015)
      Mad
      suppressible
      Dad
      (Tsuneizumi et al., 1997)
      Mov10
      suppressible
      Dad
      (Takemura et al., 2021)
      nmo
      enhanceable
      Dad
      (Zeng et al., 2007)
      pbl
      suppressible
      Dad
      (Gregory et al., 2007)
      pnr
      enhanceable
      Dad
      (Pena-Rangel et al., 2002)
      RhoGAP92B
      suppressible
      Dad
      (Nahm et al., 2010)
      shn
      enhanceable
      Dad
      (Toraason et al., 2024)
      Smn
      suppressible
      Dad
      (Chang et al., 2008)
      tkv
      suppressible
      Dad
      (Tsuneizumi et al., 1997)
      upd1
      enhanceable
      Dad
      (Bach et al., 2003)
      External Data
      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)
      3R
      Recombination map
      3-59
      Cytogenetic map
      89E11-89E11
      Sequence location
      FlyBase Computed Cytological Location
      Cytogenetic map
      Evidence for location
      89E11-89E11
      Limits computationally determined from genome sequence between P{EP}CG5208EP562&P{EP}DadEP3196 and P{PZ}l(3)0788207882
      Experimentally Determined Cytological Location
      Cytogenetic map
      Notes
      References
      89E-89E
      (Tsuneizumi, 1997.5.21)
      89E10-89E11
      (determined by in situ hybridisation)
      (Spradling et al., 1999, BDGP Project Members, 1994-1999)
      89E-89F
      (determined by in situ hybridisation)
      (Tsuneizumi et al., 1997)
      Experimentally Determined Recombination Data
      Location

      3-59

      (FlyBase, 2016)

      3-59.0

      (Comeron et al., 2012)
      Left of (cM)
      Right of (cM)
      Notes
      Stocks and Reagents
      Stocks (32)
      Genomic Clones (18)
       

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

      cDNA Clones (65)
       

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

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

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

      cDNA Clones, End Sequenced (ESTs)
      BDGP DGC clones
      Other clones
      RNAi and Array Information
      Linkouts
      DRSC - Results frm RNAi screens
      Antibody Information
      Laboratory Generated Antibodies
       
      Commercially Available Antibodies
       
      Cell Line Information
      Publicly Available Cell Lines
       
        Other Stable Cell Lines
         
          Other Comments

          Area matching Drosophila DAD gene, Acc. No. AB004232.

          (Davies, 2001.3.15)

          Dad may be capable of antagonizing signals from both the Dpp/BMP and TGF-beta/Activin TGF-beta subfamilies.

          (Marquez et al., 2001)

          Overexpression of Dad, Hsap\MADH6 or Hsap\MADH7 have comparable, truncated leg phenotypes.

          (Marquez et al., 2001)

          Dad mutations have no effect on salivary gland formation.

          (Henderson et al., 1999)

          Loss of function alleles of Dad, enhance the CycEJP mutant eye phenotype.

          (Horsfield et al., 1998)

          Transcription of Dad is induced by the dpp signal. The Dad product shows weak homology to Mad.

          (Tsuneizumi and Tabata, 1997)

          The pattern organizing mechanism governed by dpp may involve a negative feedback circuit mediated by Dad.

          (Tsuneizumi and Tabata, 1997)

          The transcription of Dad is induced by dpp, while the action of Dad antagonizes dpp.

          (Tsuneizumi et al., 1997)
          Relationship to Other Genes
          Source for database merge of

          Source for merge of: Dad anon- EST:fe1F5

          (FlyBase, 1996-)

          Source for merge of: Dad l(3)j1E4

          (Henderson et al., 1999)

          Source for merge of: Dad EP3196

          (Bidet et al., 2003)
          Additional comments

          Dad can antagonize BMP signalling in Xenopus embryos.

          (Tsuneizumi et al., 1997)
          Nomenclature History
          Source for database identify of
          Nomenclature comments
          Etymology
          Synonyms and Secondary IDs (17)
          Reported As
          Symbol Synonym
          1883
          (Tsuneizumi and Tabata, 1997, Tsuneizumi et al., 1997)
          CG5201
          (National Institute of Genetics Fly Stocks, 2023-, Hassan et al., 2018, Hosono et al., 2015, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Perkins et al., 2009.3.31, Quinones-Coello, 2007, Mohit et al., 2006, Molnar et al., 2006)
          DAD
          (Nguyen et al., 2016, Lembong et al., 2009, Tsuneizumi, 1997.5.21)
          Dad
          (Bandyadka et al., 2025, Mohar et al., 2025, Tesson and Vincent, 2025, Beadle et al., 2024, Li et al., 2024, Wang et al., 2024, Delbare et al., 2023, Liu et al., 2023, Olivares et al., 2023, Prasad et al., 2023, Yang et al., 2023, Akiyama et al., 2022, Gera et al., 2022, Kharrat et al., 2022, Montanari et al., 2022, Anqueira-González et al., 2021, Kramer et al., 2021, Sharifkhodaei and Auld, 2021, Shidlovskii et al., 2021, Takemura et al., 2021, FlyBase Genome Annotators, 2020, Fujisawa et al., 2020, Gou et al., 2020, Lamiré et al., 2020, Li et al., 2020, Thestrup et al., 2020, Zhang and Cai, 2020, Angulo et al., 2019, Chang et al., 2019, FlyBase Genome Annotators, 2019-, Gao et al., 2019, Gui et al., 2019, Hopkins et al., 2019, Houtz et al., 2019, Leiblich et al., 2019, Li et al., 2019, Svendsen et al., 2019, Tian et al., 2019, Yee et al., 2019, Chandran et al., 2018, Chen et al., 2018, Hassan et al., 2018, Kwon et al., 2018, Stephano et al., 2018, Tseng et al., 2018, Yeung et al., 2018, Beer and Wehman, 2017, Chen et al., 2017, Erceg et al., 2017, Jeibmann et al., 2017, Li et al., 2017, Liu and Jin, 2017, Takemura and Nakato, 2017, Transgenic RNAi Project members, 2017-, Barton et al., 2016, Djabrayan and Casanova, 2016, Sharifkhodaei et al., 2016, Beich-Frandsen et al., 2015, Bieli et al., 2015, Carrillo et al., 2015, Charbonnier et al., 2015, Dolezal et al., 2015, Gene Disruption Project members, 2015-, Golovnin et al., 2015, Lu et al., 2015, Wang et al., 2015, West et al., 2015, Zhou et al., 2015, Ashwal-Fluss et al., 2014, Corrigan et al., 2014, Roy et al., 2014, Tseng et al., 2014, Bausek, 2013, Chang et al., 2013, Guo et al., 2013, Jin et al., 2013, Li et al., 2013, Li et al., 2013, Mbodj et al., 2013, Paul et al., 2013, Peterson and O'Connor, 2013, Saunders et al., 2013, Umulis and Othmer, 2013, Weavers and Skaer, 2013, Xin et al., 2013, Xu et al., 2013, Zhu et al., 2013, Japanese National Institute of Genetics, 2012.5.21, Kagey et al., 2012, Leiblich et al., 2012, Lu et al., 2012, Raftery and Umulis, 2012, Spokony and White, 2012.11.14, Awofala, 2011, Dworkin et al., 2011, Eliazer and Buszczak, 2011, Galindo et al., 2011, Harris and Ashe, 2011, Kirilly et al., 2011, König et al., 2011, Liu et al., 2011, Ogiso et al., 2011, Wang et al., 2011, Ables and Drummond-Barbosa, 2010, Ballard et al., 2010, Chen et al., 2010, Firth et al., 2010, Leatherman and Dinardo, 2010, Li et al., 2010, Sato et al., 2010, Guo and Wang, 2009, Jaramillo et al., 2009, Umemori et al., 2009, Widmann and Dahmann, 2009, Ziv et al., 2009, Chang et al., 2008, Franciscovich et al., 2008, Jin et al., 2008, Kamiya et al., 2008, Kucherenko et al., 2008, Ng, 2008, Pinto et al., 2008, Shen et al., 2008, Wang et al., 2008, Yakoby et al., 2008, Yakoby et al., 2008, Zahedi et al., 2008, Zartman et al., 2008, Zhao et al., 2008, Buszczak et al., 2007, Crickmore and Mann, 2007, de Navascués and Modolell, 2007, Goold and Davis, 2007, Grammont, 2007, Gregory et al., 2007, Quinones-Coello, 2007, Song et al., 2007, Tyler et al., 2007, Wada et al., 2007, Yavari et al., 2007, Zeitlinger et al., 2007, Zeng et al., 2007, Chen and Schupbach, 2006, Choksi et al., 2006, Dudu, 2006, Dudu et al., 2006, Dworkin and Gibson, 2006, Molnar et al., 2006, Narbonne-Reveau et al., 2006, Payne and Braun, 2006, Stultz et al., 2006, Ward et al., 2006, Jordan et al., 2005, Kirilly et al., 2005, Kruger, 2005, Rogulja and Irvine, 2005, Xi and Xie, 2005, Bunt and Hime, 2004, Martin-Castellanos and Edgar, 2002, Xie and Spradling, 1998)
          EP(3)3196
          (Pena-Rangel et al., 2002)
          EP3196
          Smad7
          (van Soldt and Cardoso, 2020)
          anon-EST:fe1F5
          dad
          (Ell et al., 2024, Ju et al., 2024, Raicu et al., 2024, Matsuda and Affolter, 2023, Fan et al., 2021, Bakker et al., 2020, Grandon et al., 2019, Huang et al., 2019, Vuilleumier et al., 2019, Eusebio et al., 2018, Kanai et al., 2018, Powers and Srivastava, 2018, Setiawan et al., 2018, Heo et al., 2017, Li et al., 2017, Revaitis et al., 2017, Zhang et al., 2017, Deshpande et al., 2016, Huang et al., 2016, Lee et al., 2016, Testa and Dworkin, 2016, Wang et al., 2016, Yu et al., 2016, Ayyaz et al., 2015, Bieli et al., 2015, Denes et al., 2015, Fried and Iber, 2015, Liu et al., 2015, Menon et al., 2015, Zhao et al., 2015, Liu et al., 2014, Marmion et al., 2013, Nahm et al., 2013, Shi et al., 2013, Boulanger et al., 2012, Fuentes-Medel et al., 2012, Hadar et al., 2012, Pennetier et al., 2012, Ziv et al., 2012, Goldstein et al., 2011, Hamaratoglu et al., 2011, Kaneko et al., 2011, Ogiso et al., 2011, Oh and Irvine, 2011, Rodal et al., 2011, Szuperák et al., 2011, Wartlick et al., 2011, Ballard et al., 2010, Bunt et al., 2010, Leatherman and Dinardo, 2010, Nahm et al., 2010, Ninov et al., 2010, Vuilleumier et al., 2010, Weiss et al., 2010, Xia et al., 2010, Cruz et al., 2009, Foronda et al., 2009, Hayashi et al., 2009, Liu et al., 2009, McKay et al., 2009, Umemori et al., 2009, Chang et al., 2008, Chau et al., 2008, Franciscovich et al., 2008, Jiang et al., 2008, O'Connor-Giles et al., 2008, Shen et al., 2008, Barrio et al., 2007, Lindner et al., 2007, O'Keefe et al., 2007, Shravage et al., 2007, Tanaka et al., 2007, Umemori et al., 2007, Weiss et al., 2007, Xu et al., 2007, Dworkin and Gibson., 2006, Mohit et al., 2006, Molnar et al., 2006, Narbonne-Reveau et al., 2006, Weiss et al., 2006, Eaton and Davis, 2005, Sotillos and De Celis, 2005, Gilboa and Lehmann, 2004, Shivdasani and Ingham, 2003, Adachi-Yamada and O'Connor, 2002, Cavodeassi et al., 2002, Nakagoshi et al., 2002, Ribeiro et al., 2002, Sanyal and Ramaswami, 2002, Sweeney and Davis, 2002, Kubota et al., 2000, Kyoda et al., 2000, Marty et al., 2000, Kudoh, 1999, Kudoh, 1999, Podos and Ferguson, 1999, Cho and Blitz, 1998, Henderson and Andrew, 1998, Horsfield et al., 1998, Whitman, 1998)
          l(3)j1E4
          (Johannes and Preiss, 2002)
          Name Synonyms
          Daughter against dpp
          (Wang et al., 2016)
          Daughters against DPP
          (Lembong et al., 2009)
          Daughters against Dpp
          (Charbonnier et al., 2015, Roy et al., 2014, Bausek, 2013, Harris and Ashe, 2011, Umemori et al., 2009, Pinto et al., 2008, Tanaka et al., 2007)
          Daughters against dpp
          (Zhang and Cai, 2020, Takemura and Nakato, 2017, Djabrayan and Casanova, 2016, Xin et al., 2013, Eliazer and Buszczak, 2011, Guo and Wang, 2009, Chang et al., 2008, Gregory et al., 2007, Dworkin and Gibson, 2006, Jordan et al., 2005, Li and White, 2003)
          anon-fast-evolving-1F5
          daughter against dpp
          (Jiang et al., 2008)
          daughters against dpp
          (Yu et al., 2016, Bieli et al., 2015, Ogiso et al., 2011, Oh and Irvine, 2011, Vuilleumier et al., 2010, Foronda et al., 2009, Zhao et al., 2008, Lindner et al., 2007, Umemori et al., 2007, Tsuneizumi, 1997.5.21)
          Secondary FlyBase IDs
          • FBgn0024521
          • FBgn0025250
          • FBgn0062662
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 74 )
          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/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
          Other crossreferences
          AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
          DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
          EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
          FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
          FlyMine - An integrated database for Drosophila genomics
          InterPro - A database of protein families, domains and functional sites
          KEGG Genes - Molecular building blocks of life in the genomic space.
          MARRVEL_MODEL - MARRVEL (model organism gene)
          Linkouts
          BioGRID - A database of protein and genetic interactions.
          Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
          DroID - A comprehensive database of gene and protein interactions.
          DRSC - Results frm RNAi screens
          Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
          FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
          FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
          Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
          Flygut - An atlas of the Drosophila adult midgut
          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 (428)