FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\put
Open Close
General Information
Symbol
Dmel\put
Species
D. melanogaster
Name
punt
Annotation Symbol
CG7904
Feature Type
FlyBase ID
FBgn0003169
Gene Model Status
Stock Availability
Enzyme Name (EC)
receptor protein serine/threonine kinase (2.7.11.30)
Gene Summary
punt (put) encodes a transforming growth factor beta type II receptor that functions in both Dpp/BMP and Activin signaling. Pathway specificity in signaling output is determined by which type I receptor (Dpp/BMP or Activin one) is engaged in the complex with the product of put. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
Also Known As

Atr-II, l(3)10460, TGF-β

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
3-55
RefSeq locus
NT_033777 REGION:14619030..14625703
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (35 terms)
Molecular Function (8 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
Biological Process (22 terms)
Terms Based on Experimental Evidence (20 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:wit; FB:FBgn0024179
inferred from physical interaction with FLYBASE:babo; FB:FBgn0011300
involved_in axon guidance
inferred from mutant phenotype
inferred from mutant phenotype
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
involved_in head involution
inferred from mutant phenotype
inferred from mutant phenotype
involved_in neuron development
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
Cellular Component (5 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
located_in membrane
inferred from electronic annotation with InterPro:IPR000333, InterPro:IPR000472
located_in plasma membrane
inferred from sequence model
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the protein kinase superfamily. TKL Ser/Thr protein kinase family. TGFB receptor subfamily. (Q24468)
Catalytic Activity (EC/Rhea)
transmembrane receptor protein serine/threonine kinase activity
(1) L-seryl-[receptor-protein] + ATP = O-phospho-L-seryl-[receptor- protein] + ADP + H(+) (2.7.11.30)
(2) L-threonyl-[receptor-protein] + ATP = O-phospho-L-threonyl-[receptor- protein] + ADP + H(+) (2.7.11.30)
RHEA 18673:
Summaries
Gene Snapshot
punt (put) encodes a transforming growth factor beta type II receptor that functions in both Dpp/BMP and Activin signaling. Pathway specificity in signaling output is determined by which type I receptor (Dpp/BMP or Activin one) is engaged in the complex with the product of put. [Date last reviewed: 2019-03-14]
Pathway (FlyBase)
ACTIVIN SIGNALING PATHWAY CORE COMPONENTS -
The activin signaling pathway is one of two branches of Transforming Growth Factor-β family signaling in Drosophila. The binding of an activin family dimer to a heterodimeric serine/threonine kinase receptor complex results in the phosphorylation of Smox, a members of the Smad family. Smox forms a complex with the co-Smad, Med. This complex translocates into the nucleus and regulates the transcription of target genes in concert with other nuclear cofactors. (Adapted from FBrf0236482.)
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.)
Gene Group (FlyBase)
TGF-BETA TYPE II RECEPTORS -
The Transforming Growth Factor-β Receptor (TGF-βR) family are transmembrane serine/threonine kinases. They form heterodimeric receptor complexes composed of type I and type II subunits. Upon ligand binding, the constitutively active type II receptor phosphorylates the type I receptor which, in turn, phosphorylates a member of the Smad family. (Adapted from FBrf0236482.)
Protein Function (UniProtKB)
Type II transmembrane receptor serine/threonine kinase involved in activin-like TGF-beta signaling and dpp/BMP-like signaling pathways (PubMed:19766717, PubMed:22563507, PubMed:7697719, PubMed:7697720, PubMed:8289834, PubMed:8415726). Forms a complex with type I transmembrane receptor serine/threonine kinases, and upon activation by signaling ligands induces the type I receptor to phosphorylate downstream receptor-regulated SMAD (R-SMAD) transcription factors, inducing their nuclear localization and resulting in regulation of ligand-responsive genes (PubMed:17119022, PubMed:18820452, PubMed:7697720, PubMed:8289834). Receptor for the activin-like TGF-beta ligands daw and Actbeta/activin when in complex with the type I receptor babo (PubMed:10320478, PubMed:17119022, PubMed:18820452, PubMed:8289834). Activation by daw requires babo isoform C (PubMed:19766717). Receptor for the BMP-like ligand dpp when associated with the type I receptors tkv or sax (PubMed:7697719, PubMed:7697720). Activation by activin-like TGF-beta ligands, including daw and Actbeta/activin, results in phosphorylation of the R-SMAD Smox/SMAD2 (PubMed:10320478, PubMed:17119022, PubMed:18820452, PubMed:19766717, PubMed:8289834). Activation by BMP-like ligands, including gbb and dpp, results in phosphorylation of the R-SMAD Mad (PubMed:18820452, PubMed:19766717). Involved in dpp-signaling-dependent dorsoventral patterning of the epidermis during embryogenesis (PubMed:7697719, PubMed:7697720, PubMed:9504926). Involved in axonal and dendritic pruning and neural remodeling of mushroom body gamma neurons during metamorphosis, probably through regulating expression of the ecdysteroid receptor EcR; functions redundantly with wit (PubMed:12581521). Involved in dpp/BMP-like signaling during development of the embryonic gut tracheal system (PubMed:7697719, PubMed:7697720). Required for dpp/BMP-like signaling involved in imaginal disc development (PubMed:9504926). Involved in differentiation of dorsal cluster and ellipsoid body adult-specific neurons prior to metamorphosis (PubMed:16437159). Required in motoneurons for axon guidance or outgrowth of ISNb and SNa axons (PubMed:17119022). Required in somatic cyst cells for regulation of germ cell proliferation in testis (PubMed:9334286). Involved in adult wing development and wing-vein formation (PubMed:7697720).
(UniProt, Q24468)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
put: punt
Homozygous lethal in embryo. Dorsal part of embryo open.
Summary (Interactive Fly)

receptor for Dpp - type II TGFß receptor - functions in both Dpp/BMP and Activin signaling - punt mutants lack Ultrabithorax and dpp expression in the visceral mesoderm and fail to induce labial in the adjacent endodermal cells - pathway specificity in signaling output is determined by which type I receptor (Dpp/BMP or Activin one) is engaged in the complex with Put

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

Please see the JBrowse view of Dmel\put 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 Q24468)

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

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

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Gene model reviewed during 5.44

Stop-codon suppression (UAG) postulated; FBrf0216885.

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

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)
FBtr0082977
2713
516
FBtr0330227
2713
520
FBtr0334030
4013
516
Additional Transcript Data and Comments
Reported size (kB)

2.7 (unknown)

2.7 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0082436
58.6
516
6.74
FBpp0306147
58.6
516
6.74
Polypeptides with Identical Sequences

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

516 aa isoforms: put-PA, put-PC
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

Interacts with babo; forms a heteromeric receptor complex that can bind the activin-like ligand Actbeta (PubMed:8289834). As part of the babo-put complex associates with Smox; dissociates upon phosphorylation of Smox (PubMed:10320478, PubMed:9887103). Interacts with tkv; forms a heteromeric receptor complex that can bind BMP-like ligands (PubMed:7697720). Interacts with sax; forms a heteromeric receptor complex that can bind BMP-like ligands (PubMed:7697720).

(UniProt, Q24468)
Domain

The basolateral targeting (BLT) motif within the cytoplasmic region determines basolateral membrane localization in wing disc epithelial cells.

(UniProt, Q24468)
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\put using the Feature Mapper tool.

External Data
Crossreferences
Linkouts
Expression Data
Testis-specificity index

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

-1.25

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

put transcripts are detected throughout development on northern blots. They are abundant in nurse cells and in the developing oocyte. The transcript persists at high levels in the embryo until cellularization. Transcript levels decrease at cellularization but remain highest under the pole cells. At the start of gastrulation, transcripts are observed in the invaginating mesoderm. By stage 11, expression is observed along the entire length of the germ band in the mesoderm as well as in the developing gut. Expression continues in the fore- mid- and hindgut throughout most of embryogenesis. Third instar larval imaginal discs show prominant put expression.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

put protein is observed throughout the embryo at syncytial blastoderm stage. At the start of gastrulation, Pburs protein is observed in the invaginating mesoderm. By stage 11 it is expressed along the entire length of the germ band in the mesoderm as well as in the developing gut. Expression continues in the fore- mid- and hindgut throughout most of embryogenesis.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\put 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
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 25 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 46 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of put
Transgenic constructs containing regulatory region of put
Aberrations (Deficiencies and Duplications) ( 7 )
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
dorsal cluster neuron & axon | somatic clone
dorsal cluster neuron & neurite | somatic clone
dorsal cluster neuron & neurite | somatic clone | conditional ts
scutum & macrochaeta, with Scer\GAL4hs.PB
scutum & macrochaeta, with Scer\GAL4tsh-Gal4-1
sensory mother cell & dorsal mesothoracic disc
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (37)
12 of 14
Yes
Yes
11 of 14
No
Yes
1  
4 of 14
No
Yes
3 of 14
No
No
3 of 14
No
Yes
2 of 14
No
No
3  
2 of 14
No
No
1 of 14
No
No
0  
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
4  
1 of 14
No
No
3  
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
2  
1 of 14
No
No
1 of 14
No
No
3  
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
9  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (25)
12 of 14
Yes
Yes
11 of 14
No
Yes
4 of 14
No
Yes
3 of 14
No
No
3 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
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Mus musculus (laboratory mouse) (24)
12 of 14
Yes
Yes
11 of 14
No
Yes
4 of 14
No
Yes
3 of 14
No
Yes
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
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (21)
9 of 13
Yes
Yes
6 of 13
No
Yes
4 of 13
No
No
2 of 13
No
No
2 of 13
No
Yes
2 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
Danio rerio (Zebrafish) (36)
12 of 14
Yes
Yes
10 of 14
No
Yes
10 of 14
No
Yes
10 of 14
No
Yes
4 of 14
No
Yes
3 of 14
No
No
3 of 14
No
Yes
3 of 14
No
Yes
2 of 14
No
No
1 of 14
No
Yes
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (16)
9 of 14
Yes
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 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) (16)
11 of 12
Yes
Yes
1 of 12
No
No
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (122)
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
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Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
No
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Yes
No
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Yes
No
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Yes
No
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Yes
No
1 of 13
Yes
No
1 of 13
Yes
Yes
1 of 13
Yes
No
1 of 13
Yes
Yes
1 of 13
Yes
No
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Yes
No
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Yes
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Yes
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Yes
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No
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (1)
1 of 13
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:put. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (15)
7 of 13
3 of 13
3 of 13
3 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 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 ( 1 )
Allele
Disease
Evidence
References
Potential Models Based on Orthology ( 5 )
Modifiers Based on Experimental Evidence ( 0 )
Allele
Disease
Interaction
References
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.
Interactions
Summary of Physical Interactions
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
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
enhanceable
External Data
Subunit Structure (UniProtKB)
Interacts with babo; forms a heteromeric receptor complex that can bind the activin-like ligand Actbeta (PubMed:8289834). As part of the babo-put complex associates with Smox; dissociates upon phosphorylation of Smox (PubMed:10320478, PubMed:9887103). Interacts with tkv; forms a heteromeric receptor complex that can bind BMP-like ligands (PubMed:7697720). Interacts with sax; forms a heteromeric receptor complex that can bind BMP-like ligands (PubMed:7697720).
(UniProt, Q24468 )
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.
SignaLink - A signaling pathway resource with multi-layered regulatory networks.
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3R
Recombination map
3-55
Cytogenetic map
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
88C9-88C9
Limits computationally determined from genome sequence between P{PZ}cv-c06951 and P{PZ}put10460
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
88C9-88C10
(determined by in situ hybridisation)
88D-88D
(determined by in situ hybridisation)
Experimentally Determined Recombination Data
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (27)
Genomic Clones (26)
cDNA Clones (63)
 

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

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

        put is required during the late larval stage for development of adult-specific neurons.

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

        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.

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

        The embryonic dorsoventral patterning and dorsal closure phenotypes seen in put mutants correspond to distinct maternal and zygotic requirements for put.

        shn and put are required to limit transient amplification of germ cells. Mosaic analysis demonstrates shn and put act within somatic cyst cells that surround germ cells, rather than in germ cells. Thus a cyst-cell-derived signal restricts germ cell proliferation and this signal is initiated by input from a member of the TGF-β superfamily. Thus, a signal relay regulates progression through the germline stem cell lineage.

        dpp receptors tkv and put are not required for photoreceptor cell differentiation. tkv and put play a nonessential role in morphogenetic furrow progression, but are required for initiation of the furrow at the posterior margin. Ectopic activation of the dpp pathway does not lead to ectopic neuronal differentiation.

        dpp receptors put and tkv, or the shn transcription factor, are autonomously required for cell proliferation in the entire developing wing. The dpp signal has to travel several cell diameters from its source in order to reach all cells that require its signal.

        put gene product is a dpp type II receptor, type II receptors have a dual ligand binding specificity.

        Genetic interactions and the strikingly similar phenotypes in shn and dpp receptors encoded by thv and put suggest the shn plays a downstream role in dpp signalling.

        Activin type II receptor is encoded by the put gene. Loss of the maternal function of the put gene disrupts a signalling pathway required for embryonic pattern function, causing cells throughout the embryo to adopt a fate normally reserved for those at the ventral surface.

        Maternal and zygotic loss of put ventralises the embryo.

        Mitotic recombination analysis demonstrates a requirement for maternal put gene product in dorsal-ventral patterning.

        The P-element line 10460 fails to complement the EMS allele put135.

        put and tkv encode type II and type I receptors, both act in concert to mediate all known dpp signalling processes in the developing midgut.

        put encodes a type II receptor for dpp, Rec88D.

        Ligand binding studies reveal put gene product binds activin with high specificity. The sequence and binding characteristics of put indicate that only a small number of amino acids are required for specification of activin binding.

        TGF-β like activity is present in cell extracts.

        put mutants display failure of dorsal closure.

        Relationship to Other Genes
        Source for database merge of
        Additional comments
        Nomenclature History
        Source for database identify of

        Source for identity of: put CG7904

        Nomenclature comments
        Etymology
        Synonyms and Secondary IDs (26)
        Reported As
        Symbol Synonym
        Act-r
        Atr
        Atr88CD
        AtrII
        Tgf-r
        dlhC
        l(3)j5A5
        put
        (Bischoff et al., 2025, Klimkowski Arango and Morgante, 2025, Sheng et al., 2025, Ell et al., 2024, Sutton et al., 2024, Bastin and Eleftherianos, 2023, Bauer et al., 2023, Lassetter et al., 2023, Jiang et al., 2022, Peterson et al., 2022, Saha et al., 2022, Bilder et al., 2021, Everetts et al., 2021, Herrera et al., 2021, Hertenstein et al., 2021, Lebo et al., 2021, Ostalé et al., 2021, Hinnant et al., 2020, Port et al., 2020, Wei et al., 2020, Bangi et al., 2019, Hall et al., 2019, Ng, 2019.8.30, Story et al., 2019, Wang et al., 2019, Xu et al., 2019, Ameku et al., 2018, Gene Disruption Project members, 2018-, Hoyer et al., 2018, Le et al., 2018, Song et al., 2018, Wang et al., 2018, Yeung et al., 2018, Houtz et al., 2017, Jordán-Álvarez et al., 2017, Ma et al., 2017, Song et al., 2017, Transgenic RNAi Project members, 2017-, Moulton and Letsou, 2016, Sarov et al., 2016, Sulkowski et al., 2016, Aradska et al., 2015, Ugrankar et al., 2015, Ashwal-Fluss et al., 2014, Roy et al., 2014, Sopko et al., 2014, Sulkowski et al., 2014, Tian and Jiang, 2014, Curtis et al., 2013, Hevia and de Celis, 2013, Li et al., 2013, Marinho et al., 2013, Saunders et al., 2013, Le and Wharton, 2012, Jungreis, 2011.11.18, Michel et al., 2011, Neumüller et al., 2011, Yuva-Aydemir et al., 2011, Ables and Drummond-Barbosa, 2010, Salzer et al., 2010, Cook et al., 2009.11.25, Wagner et al., 2009, Yan et al., 2009, Ng, 2008, Wang et al., 2008, Adryan et al., 2007, Chen et al., 2007, Cordero et al., 2007, Dietzl et al., 2007, Fernández et al., 2007, Zeitlinger et al., 2007, Anderson et al., 2006, Bangi and Wharton, 2006, Joulia et al., 2006, Li and Li, 2006, Roote, 2006.8.30, Xie et al., 2005, Bennett and Alphey, 2002, Pickeral et al., 2000, Riesgo-Escovar and Hafen, 1997)
        Name Synonyms
        Activin-A-receptor-88CD
        DPP receptor
        TGF-β-type-receptor-like
        activin receptor
        activin receptor type II
        dorsal holes C
        Secondary FlyBase IDs
        • FBgn0000467
        • FBgn0004912
        • FBgn0004920
        • FBgn0010942
        • FBgn0011375
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 61 )
        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)
        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
        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
        SignaLink - A signaling pathway resource with multi-layered regulatory networks.
        References (445)