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General Information
Symbol
Dmel\sav
Species
D. melanogaster
Name
salvador
Annotation Symbol
CG33193
Feature Type
FlyBase ID
FBgn0053193
Gene Model Status
Stock Availability
Gene Snapshot
salvador (sav) encodes a scaffold protein involved in the Hippo signalling pathway. It promotes membrane recruitment of the kinase encoded by hpo and its assembly with the product of wts. [Date last reviewed: 2018-11-15]
Also Known As
shar-pei
Key Links
Genomic Location
Cytogenetic map
Sequence location
3R:23,058,609..23,061,304 [-]
Recombination map
3-77
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
Predictions / Assertions
-
Summaries
Pathway (FlyBase)
Hippo Signaling Pathway Core Components -
The Hippo signaling pathway is an intracellular kinase cascade in which hpo kinase in complex with sav, phosphorylates wts kinase which, in turn, phosphorylates yki transcriptional co-activator leading to its cytosolic retention. Activation of the Hippo pathway results in the down-regulation of cell proliferation and up-regulation of apoptosis, limiting tissue size. (Adapted from FBrf0224870).
Protein Function (UniProtKB)
Plays a key role in the Hippo/SWH (Sav/Wts/Hpo) signaling pathway, a signaling pathway that plays a pivotal role in organ size control and tumor suppression by restricting proliferation and promoting apoptosis. The core of this pathway is composed of a kinase cascade wherein Hippo (Hpo), in complex with its regulatory protein Salvador (Sav), phosphorylates and activates Warts (Wts) in complex with its regulatory protein Mats, which in turn phosphorylates and inactivates the Yorkie (Yki) oncoprotein. The Hippo/SWH signaling pathway inhibits the activity of the transcriptional complex formed by Scalloped (sd) and Yki and the target genes of this pathway include cyclin-E (cycE), diap1 and bantam. Required for cell cycle exit in eye imaginal disk and hid-induced apoptotic cell deaths that are part of normal retinal development. Activation of Drice in eye imaginal disk by either Hid or Rpr is almost completely blocked by Sav expression.
(UniProt, Q9VCR6)
Summary (Interactive Fly)
scaffold protein - restricts cell numbers in vivo by functioning as a dual regulator of cell proliferation and apoptosis - physically interacts with Warts - the stability of Salvador (Sav), which is believed to promote Hippo/Warts association, is crucially dependent on its binding partner Hippo
Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
2

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

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

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Comments on Gene Model
Gene model reviewed during 5.48
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0084356
2235
608
FBtr0303509
2220
603
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
FBpp0083749
68.1
608
9.92
FBpp0292543
67.5
603
9.90
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)
Interacts with Wts via its WW domains. Interacts (via FBM motif) with Mer (via FERM domain). Interacts with Kibra. Interacts with Hpo (via SARAH domain). Interacts with jub.
(UniProt, Q9VCR6)
Post Translational Modification
Phosphorylated by Hpo.
(UniProt, Q9VCR6)
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\sav 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 (23 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from physical interaction with UniProtKB:Q9VFG8
(assigned by UniProt )
inferred from physical interaction with UniProtKB:Q9VY77
(assigned by UniProt )
Terms Based on Predictions or Assertions (0 terms)
Biological Process (16 terms)
Terms Based on Experimental Evidence (12 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from direct assay
inferred from genetic interaction with FLYBASE:wts; FB:FBgn0011739
inferred from genetic interaction with FLYBASE:wts; FB:FBgn0011739, FLYBASE:sav; FB:FBgn0053193
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:hid; FB:FBgn0003997
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN002569181
(assigned by GO_Central )
traceable author statement
(assigned by UniProt )
inferred from biological aspect of ancestor with PANTHER:PTN000817101
(assigned by GO_Central )
Cellular Component (5 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
inferred from direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000817101
(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
inferred from direct assay
inferred from direct assay
inferred from direct assay
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\sav 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 ( 14 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 20 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of sav
Transgenic constructs containing regulatory region of sav
Deletions and Duplications ( 3 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
dorsal multidendritic neuron ddaC | somatic clone & dendritic tree
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
9 of 15
Yes
Yes
1 of 15
No
Yes
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
8 of 15
Yes
Yes
1 of 15
No
Yes
Rattus norvegicus (Norway rat) (2)
8 of 13
Yes
Yes
1 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (1)
6 of 12
Yes
Yes
Danio rerio (Zebrafish) (2)
7 of 15
Yes
Yes
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (1)
8 of 15
Yes
Yes
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG0919070C )
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 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) ( EOG091502GM )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
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) ( EOG090W094A )
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
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X090H )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Strigamia maritima
European centipede
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) ( EOG091G09KI )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (2)
2 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 ( 0 )
Allele
Disease
Evidence
References
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 1 )
Allele
Disease
Interaction
References
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
suppressible
External Data
Subunit Structure (UniProtKB)
Interacts with Wts via its WW domains. Interacts (via FBM motif) with Mer (via FERM domain). Interacts with Kibra. Interacts with Hpo (via SARAH domain). Interacts with jub.
(UniProt, Q9VCR6 )
Linkouts
BioGRID - A database of protein and genetic interactions.
DroID - A comprehensive database of gene and protein interactions.
InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Signaling Pathways (FlyBase)
Hippo Signaling Pathway Core Components -
The Hippo signaling pathway is an intracellular kinase cascade in which hpo kinase in complex with sav, phosphorylates wts kinase which, in turn, phosphorylates yki transcriptional co-activator leading to its cytosolic retention. Activation of the Hippo pathway results in the down-regulation of cell proliferation and up-regulation of apoptosis, limiting tissue size. (Adapted from FBrf0224870).
Metabolic Pathways
External Data
Linkouts
KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3R
Recombination map
3-77
Cytogenetic map
Sequence location
3R:23,058,609..23,061,304 [-]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
94D10-94D10
Limits computationally determined from genome sequence between P{lacW}GclmL0580 and P{EP}hhEP3521
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
Experimentally Determined Recombination Data
Notes
Located on the right arm of chromosome 3
Stocks and Reagents
Stocks (8)
Genomic Clones (16)
 

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

cDNA Clones (118)
 

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

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

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

    cDNA Clones, End Sequenced (ESTs)
    BDGP DGC clones
    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
     
    polyclonal
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for database merge of
    Source for merge of: salvador CG13831
    Source for merge of: CG13831 CG13832
    Source for merge of: sav anon-WO0172774.152
    Additional comments
    Annotations CG13831, CG13832 merged as CG33193 in release 3 of the genome annotation.
    Source for merge of CG13831 CG13832 was a shared cDNA ( date:010720 ).
    Source for merge of sav anon-WO0172774.152 was sequence comparison ( date:051113 ).
    Other Comments
    sav acts cell autonomously in class IV neurons.
    dsRNA has been made from templates generated with primers directed against this gene.
    sav is required for the proper termination of cell proliferation during imaginal disc development.
    sav promotes both cell cycle exit and apoptosis.
    Identification: Isolated as a mutation that causes overgrowth in homozygous clones in the eye.
    Mutations in sav abolish developmentally controlled programmed cell death in homozygous clones in the eye.
    Origin and Etymology
    Discoverer
    Etymology
    The gene is called "shar-pei" because of the mutant folded cuticle phenotype in the head, which resembles the folded skin of Shar-pei dogs.
    Identification
    External Crossreferences and Linkouts ( 52 )
    Sequence Crossreferences
    NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
    GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
    GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
    RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
    UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
    UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
    Other crossreferences
    BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
    Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
    Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
    Flygut - An atlas of the Drosophila adult midgut
    Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    InterPro - A database of protein families, domains and functional sites
    KEGG Genes - Molecular building blocks of life in the genomic space.
    modMine - A data warehouse for the modENCODE project
    Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
    SignaLink - A signaling pathway resource with multi-layered regulatory networks.
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    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
    Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
    Synonyms and Secondary IDs (12)
    Reported As
    Symbol Synonym
    CG13832
    Salvador/Shar-pei
    anon-WO0172774.152
    sav
    (Snigdha et al., 2019, Gou et al., 2018, Yu and Pan, 2018, Baker, 2017, Su et al., 2017, Transgenic RNAi Project members, 2017-, Jahanshahi et al., 2016, Jukam et al., 2016, Liu et al., 2016, Mao et al., 2016, Aerne et al., 2015, Dent et al., 2015, Doggett et al., 2015, Dong et al., 2015, Keder et al., 2015, Wang and Baker, 2015, Zaessinger et al., 2015, Chen et al., 2014, Huang and Kalderon, 2014, Huang et al., 2014, Rauskolb et al., 2014, Robbins et al., 2014, Tipping and Perrimon, 2014, Guo et al., 2013, Huang et al., 2013, Jukam et al., 2013, Jukam et al., 2013, Kwon et al., 2013, Kwon et al., 2013, Yin et al., 2013, Hafezi et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Poon et al., 2012, Verghese et al., 2012, Yue et al., 2012, Bao et al., 2011, Fernández et al., 2011, Napoletano et al., 2011, Ohsawa et al., 2011, Poon et al., 2011, Baumgartner et al., 2010, Das Thakur et al., 2010, Grzeschik et al., 2010, Milton et al., 2010, Robinson et al., 2010, Shaw et al., 2010, Wasbrough et al., 2010, Parrish et al., 2009, Rhiner et al., 2009, Dutta and Baehrecke, 2008, Sun et al., 2008, Yu et al., 2008, Zhang et al., 2008, Menut et al., 2007, Parrish et al., 2007, Polesello and Tapon, 2007, Tyler and Baker, 2007, Tyler et al., 2007, Wei et al., 2007, Bennett and Harvey, 2006, Cho et al., 2006, Colombani et al., 2006, Edgar, 2006, Emoto et al., 2006, Hamaratoglu et al., 2006, Polesello et al., 2006, Firth and Baker, 2005, Mikeladze-Dvali et al., 2005, Pantalacci, 2003)
    Secondary FlyBase IDs
    • FBgn0026180
    • FBgn0039046
    • FBgn0039047
    • FBgn0046446
    Datasets (1)
    Study focus (1)
    Experimental Role
    Project
    Project Type
    Title
    • bait_protein
    Interaction map generated by purification of Hippo pathway factors, with identification of copurifying proteins by mass spectrometry.
    References (220)