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General Information
Symbol
Dmel\Rheb
Species
D. melanogaster
Name
Ras homolog enriched in brain
Annotation Symbol
CG1081
Feature Type
FlyBase ID
FBgn0041191
Gene Model Status
Stock Availability
Gene Snapshot
Ras homolog enriched in brain (Rheb) encodes a Ras homolog that is the target of the product of gig. It activates the protein kinase encoded by Tor, leading to enlarged cell size. [Date last reviewed: 2018-11-08]
Also Known As
dRheb, Ras homologue enriched in brain
Key Links
Genomic Location
Cytogenetic map
Sequence location
3R:5,568,921..5,570,491 [+]
Recombination map
3-47.5
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the small GTPase superfamily. Rheb family. (Q9VND8)
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
Predictions / Assertions
Summaries
Gene Group (FlyBase)
RAS GTPASES -
The Ras family are members of the Ras superfamily of small GTPases. Ras proteins act in signal transduction cascades, generally the activated GTPase recruits and activates downstream effectors. Members this family are typically membrane-associated via isoprenylation of a CaaX motif in the C-terminus. (Adapted from PMID:15731001).
Pathway (FlyBase)
Insulin-like Receptor Signaling Pathway Core Components -
The Insulin-like Receptor (IR) signaling pathway in Drosophila is initiated by the binding of an insulin-like peptides to the Insulin-like receptor (InR). (Adapted from FBrf0232297, FBrf0230017 and FBrf0229989.)
Protein Function (UniProtKB)
Binds GTP and exhibits intrinsic GTPase activity (By similarity). Activates the protein kinase activity of TORC1, and thereby plays a role in the regulation of apoptosis (PubMed:22493059). Stimulates the phosphorylation of S6K through activation of TORC1 signaling (PubMed:22493059). May also have a role in activating TORC2 signaling (PubMed:22493059).
(UniProt, Q9VND8)
Summary (Interactive Fly)
Ras superfamily of G-proteins - functions downstream of the tumor suppressors Tsc1 and Tsc2 (Gigas in Drosophila), with Tsc2 functioning as a GAP for Rheb - signals to the growth regulator known as Target of rapamycin (TOR)
Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\Rheb or the JBrowse view of Dmel\Rheb for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Comments on Gene Model
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.48
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0078693
1225
182
FBtr0078694
1059
182
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0078342
20.7
182
5.32
FBpp0078343
20.7
182
5.32
Polypeptides with Identical Sequences

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

182 aa isoforms: Rheb-PA, Rheb-PB
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
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\Rheb using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (27 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
inferred from direct assay
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000631348
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000631348
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000631348
(assigned by GO_Central )
Biological Process (23 terms)
Terms Based on Experimental Evidence (22 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
(assigned by UniProt )
inferred from mutant phenotype
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
inferred from mutant phenotype
inferred from expression pattern
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN001529822
(assigned by GO_Central )
Cellular Component (1 term)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000631348
(assigned by GO_Central )
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: maternally deposited

Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Rheb in GBrowse 2
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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 21 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 10 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Rheb
Transgenic constructs containing regulatory region of Rheb
Deletions and Duplications ( 1 )
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 v7.1)
Homo sapiens (Human) (6)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
12 of 15
Yes
Yes
 
7 of 15
No
Yes
1 of 15
No
No
 
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (4)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
12 of 15
Yes
Yes
7 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
Rattus norvegicus (Norway rat) (4)
8 of 13
Yes
Yes
5 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (4)
6 of 12
Yes
Yes
6 of 12
Yes
Yes
1 of 12
No
No
1 of 12
No
No
Danio rerio (Zebrafish) (6)
13 of 15
Yes
Yes
5 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (3)
11 of 15
Yes
Yes
1 of 15
No
No
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (4)
1 of 9
Yes
No
1 of 9
Yes
No
1 of 9
Yes
No
1 of 9
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (1)
11 of 15
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (1)
11 of 12
Yes
Yes
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG09190GGJ )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG09150BZU )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0EHT )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0E4N )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0ONO )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (11)
2 of 10
2 of 10
2 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 4 )
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 1 )
Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
 
Disease Associations of Human Orthologs (via DIOPT v7.1 and OMIM)
Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
Homo sapiens (Human)
Gene name
Score
OMIM
OMIM Phenotype
DO term
Complementation?
Transgene?
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
esyN Network Diagram
Show neighbor-neighbor interactions:
Select Layout:
Legend:
Protein
RNA
Selected Interactor(s)
Interactions Browser

Please see the Physical Interaction reports below for full details
protein-protein
Physical Interaction
Assay
References
Summary of Genetic Interactions
esyN Network Diagram
esyN Network Key:
Suppression
Enhancement

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
External Data
Linkouts
DroID - A comprehensive database of gene and protein interactions.
InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Gene Group - Pathway Membership (FlyBase)
Insulin-like Receptor Signaling Pathway Core Components -
The Insulin-like Receptor (IR) signaling pathway in Drosophila is initiated by the binding of an insulin-like peptides to the Insulin-like receptor (InR). (Adapted from FBrf0232297, FBrf0230017 and FBrf0229989.)
External Data
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3R
Recombination map
3-47.5
Cytogenetic map
Sequence location
3R:5,568,921..5,570,491 [+]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
83B2-83B2
Limits computationally determined from genome sequence between P{PZ}Snr101319&P{PZ}Itp-r83A05616 and P{lacW}Atus1938
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
Experimentally Determined Recombination Data
Location
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (11)
Genomic Clones (17)
cDNA Clones (82)
 

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

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

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

cDNA Clones, End Sequenced (ESTs)
RNAi and Array Information
Linkouts
DRSC - Results frm RNAi screens
GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
Antibody Information
Laboratory Generated Antibodies
Commercially Available Antibodies
 
Other Information
Relationship to Other Genes
Source for database identify of
Source for database merge of
Source for merge of: Rheb CG1081
Additional comments
Other Comments
RNAi screen using dsRNA made from templates generated with primers directed against this gene results in chromosome misalignment on the metaphase spindle when assayed in S2 cells in the presence of Cdc27 dsRNA. This phenotype cannot be observed when the screen is performed without Cdc27 dsRNA.
dsRNA made from templates generated with primers directed against this gene results in a reduced mean cell diameter. Silencing Rheb generates a cell-expression profile similar to that of rapamycin-treated cells.
dsRNA made from templates generated with primers directed against this gene used to treat S2 cells.
dsRNA made from templates generated with primers directed against this gene promotes the phosphorylation of Akt1 (dependent on TORC2) and inhibits phosphorylation of S6k in S2 cells. In addition, knockdown of Rheb enhances insulin-stimulated Akt1 phosphorylation.
Signaling through Tor and its upstream regulators Pi3K92E/Pi3K21B and Rheb is necessary and sufficient to suppress starvation-induced autophagy in the fat body.
dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.
Levels of Rheb mRNA are rapidly induced in response to protein starvation.
In mitotic tissues, overexpression of Rheb accelerates passage through G1-S phase without affecting rates of cell division, whereas in endoreplicating tissues, overexpression increases DNA ploidy. Overexpression of Rheb can also drive cell growth in starved animals.
Origin and Etymology
Discoverer
Etymology
Named "Rheb" after the human ortholog.
Identification
External Crossreferences and Linkouts ( 50 )
Sequence Crossreferences
NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
Other crossreferences
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
KEGG Genes - Molecular building blocks of life in the genomic space.
modMine - A data warehouse for the modENCODE project
Linkouts
DPiM - Drosophila Protein interaction map
DroID - A comprehensive database of gene and protein interactions.
DRSC - Results frm RNAi screens
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
FlyMine - An integrated database for Drosophila genomics
Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Synonyms and Secondary IDs (15)
Reported As
Symbol Synonym
Dm Rheb
Rheb
(Galenza and Foley, 2019, Kim and Choi, 2019, Lee et al., 2019, Sharma et al., 2019, Zheng, 2019.3.1, Aboudhiaf et al., 2018, Ayala et al., 2018, Inoue et al., 2018, Nowak et al., 2018, Staats et al., 2018, Tang et al., 2018, Zeng et al., 2018, Kamalesh et al., 2017, Keesey et al., 2017, Li et al., 2017, Liu and Jin, 2017, Liu et al., 2017, Melani et al., 2017, Mensah et al., 2017, Murillo-Maldonado and Riesgo-Escovar, 2017, Romero-Pozuelo et al., 2017, Wen et al., 2017, Danielsen et al., 2016, David-Morrison et al., 2016, Dobson et al., 2016, Koyama and Mirth, 2016, Kuleesha et al., 2016, Le et al., 2016, Parkhitko et al., 2016, Rabinovich et al., 2016, Vinayagam et al., 2016, Aradhya et al., 2015, Aradska et al., 2015, Das and Dobens, 2015, Francis and Ghabrial, 2015, Grotewiel and Bettinger, 2015, Kuhn et al., 2015, Lin et al., 2015, Mensah et al., 2015, Nässel et al., 2015, Nie et al., 2015, Rojas-Benitez et al., 2015, Van Bortle et al., 2015, Zhang and Baehrecke, 2015, Avet-Rochex et al., 2014, Barrio et al., 2014, Deivasigamani et al., 2014, DeVorkin et al., 2014, Frost et al., 2014, Ghosh et al., 2014, Homem et al., 2014, Mulakkal et al., 2014, Zhang et al., 2014, Gordon et al., 2013, Ibar et al., 2013, Luo et al., 2013, Natarajan et al., 2013, Nowak et al., 2013, O'Farrell et al., 2013, Parisi et al., 2013, Quan et al., 2013, Shim et al., 2013, Shin et al., 2013, Sopko and Perrimon, 2013, Tran et al., 2013, Wong et al., 2013, Zirin et al., 2013, Alvarez-Ponce et al., 2012, Avet-Rochex et al., 2012, Dimitroff et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Kapuria et al., 2012, Kaun et al., 2012, Mirth and Shingleton, 2012, Pallares-Cartes et al., 2012, Pircs et al., 2012, Tokusumi et al., 2012, Wong et al., 2012, Yaniv et al., 2012, Zitserman et al., 2012, Barth et al., 2011, Diez et al., 2011, Eddison et al., 2011, Glatter et al., 2011, Lindquist et al., 2011, Murillo-Maldonado et al., 2011, Sousa-Nunes et al., 2011, Tang et al., 2011, Willecke et al., 2011, Chang and Neufeld, 2010, Daubert et al., 2010, Haberman et al., 2010, Karbowniczek et al., 2010, Kockel et al., 2010, Kong et al., 2010, Lee et al., 2010, Li et al., 2010, Li et al., 2010, Nisoli et al., 2010, Ribeiro and Dickson, 2010, Zheng and Sehgal, 2010, Chang and Neufeld, 2009, Chang and Neufeld, 2009, Doumanis et al., 2009, Nuzhdin et al., 2009, Parrish et al., 2009, Patch et al., 2009, Sims et al., 2009, Goraksha and Neufeld, 2008, Harvey et al., 2008, Juhász et al., 2008, Kim et al., 2008, Lee et al., 2008, McNeill et al., 2008, Rehmann et al., 2008, Bakal et al., 2007, Curtis et al., 2007, Findlay et al., 2007, Grewal et al., 2007, Hall et al., 2007, Hsu and Choi, 2007, Knox et al., 2007, Lee et al., 2007, Scott et al., 2007, Scott et al., 2007, Steinhilb et al., 2007, Tseng et al., 2007, Friedman and Perrimon, 2006, Hennig et al., 2006, Herranz et al., 2006, Khurana et al., 2006, Martin-Pena et al., 2006, Patel and Tamanoi, 2006, Walker et al., 2006, Wu and Brown, 2006, Wullschleger, 2006, Yang et al., 2006, Goberdhan et al., 2005, Liebl and Featherstone, 2005, Aspuria and Tamanoi, 2004, Reis and Edgar, 2004, Wheeler et al., 2004)
Name Synonyms
RAS homolog enriched in brain
Ras homolog enriched in brain ortholog
Ras homologue enriched in brain
Ras homology enriched in brain
Secondary FlyBase IDs
  • FBgn0037343
Datasets (1)
Study focus (1)
Experimental Role
Project
Project Type
Title
  • bait_protein
Interaction map generated by purification of insulin pathway factors, with identification of copurifying proteins by mass spectrometry.
References (266)