FB2026_02 , released June 18, 2026
Gene: Dmel\Sara
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General Information
Symbol
Dmel\Sara
Species
D. melanogaster
Name
Smad anchor for receptor activation
Annotation Symbol
CG15667
Feature Type
protein_coding_gene
FlyBase ID
FBgn0026369
Gene Model Status
Current
Stock Availability
8 publicly available
Gene Summary
Smad anchor for receptor activation (Sara) encodes an endosomal protein involved in asymmetric stem cell division, intestinal stem cell homeostasis and stem cell fate determination. [Date last reviewed: 2019-09-26] (FlyBase Gene Snapshot)
All Summaries
Also Known As

dSARA

Key Links
Genomic Location
Cytogenetic map
57E6-57E6
Sequence location
Recombination map
2-95
RefSeq locus
NT_033778 REGION:21481927..21487107
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 (8 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables protein phosphatase 1 binding
inferred from physical interaction with FLYBASE:flw; FB:FBgn0000711
(Bennett et al., 2006)
inferred from physical interaction with FLYBASE:Pp1-13C; FB:FBgn0003132
(Bennett et al., 2006)
inferred from physical interaction with FLYBASE:Pp1α-96A; FB:FBgn0003134
(Bennett et al., 2006)
inferred from physical interaction with FLYBASE:Pp1-87B; FB:FBgn0004103
(Bennett et al., 2006)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
enables metal ion binding
inferred from electronic annotation with InterPro:IPR000306
(InterPro Project Members, 2004-)
Biological Process (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
involved_in asymmetric stem cell division
inferred from mutant phenotype
(Montagne and Gonzalez-Gaitan, 2014)
involved_in intestinal stem cell homeostasis
inferred from mutant phenotype
(Montagne and Gonzalez-Gaitan, 2014)
involved_in stem cell fate determination
inferred from mutant phenotype
(Montagne and Gonzalez-Gaitan, 2014)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
involved_in endosomal transport
inferred from biological aspect of ancestor with PANTHER:PTN002335850
(GO Reference Genome Project, 2011-)
Cellular Component (2 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
located_in early endosome
inferred from direct assay
(Bokel et al., 2006)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in early endosome membrane
inferred from biological aspect of ancestor with PANTHER:PTN002335850
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
Smad anchor for receptor activation (Sara) encodes an endosomal protein involved in asymmetric stem cell division, intestinal stem cell homeostasis and stem cell fate determination. [Date last reviewed: 2019-09-26]
Summary (Interactive Fly)

binds the PP1c phosphatase and its regulator Sds22 - both Notch and Delta traffic through Sara endosomes - involved in signalling bias between the pIIa-pIIb sibling cells - Sara endosomes are specifically targeted to the pIIa cell during sensory organ precursor division, mediating the transport of a pool of Notch and Delta that contribute to the activation of Notch in the pIIa - Uninflatable controls the asymmetric dispatch of the Sara endosomes

Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\Sara 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 Q7K9H6)

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 5.51

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Sara-RA
FBtr0071641
NM_079990
4621
1343
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
UniProt
RefSeq ID
GenBank
Sara-PA
FBpp0071564
146.3
1343
4.40
Q7K9H6
NP_524729
AAF46725
Polypeptides with Identical Sequences

There is only one protein coding transcript and one polypeptide associated with this gene

Additional Polypeptide Data and Comments
Reported size (kDa)
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\Sara using the Feature Mapper tool.

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

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

-0.85

Transcript Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in early endosome
inferred from direct assay
(Bokel et al., 2006)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Sara 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 ( 5 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 13 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Sara
Transgenic constructs containing regulatory region of Sara
Aberrations (Deficiencies and Duplications) ( 3 )
Inferred from experimentation ( 3 )
Gene partially disrupted in
Df(2R)Sara12
(Bokel et al., 2006)
Gene disrupted in
Df(2R)48
(Johnson et al., 2004)
Df(2R)Exel6076
(Ryder, 2004.4.26)
Inferred from location ( 5 )
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
lethal, with Scer\GAL4da.G32
lethal | embryonic stage | recessive | rescuable maternal effect
lethal | larval stage | recessive
lethal | pupal stage | recessive
lethal - all die during embryonic stage | maternal effect | recessive
partially lethal - majority die
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4tin.CΔ4
Other Phenotypes
Allele
abnormal neuroanatomy | larval stage, with Sdc48/Sdc10608
decreased occurrence of cell division | adult stage
increased cell death
increased cell death (with Df(2R)PK1)
increased cell number, with Scer\GAL4NP5130
increased cell number | adult stage
some die during embryonic stage | recessive | maternal effect | recessive
visible
Phenotype manifest in
Allele
adult midgut
adult midgut (with Df(2R)48)
anterior wing margin, with Scer\GAL4vg.int2.1
double row of wing sensilla | ectopic, with Scer\GAL4vg.int2.1
enterocyte of posterior adult midgut epithelium | somatic clone | increased number
enteroendocrine cell of posterior adult midgut epithelium | somatic clone
intestinal stem cell of posterior adult midgut epithelium, with Scer\GAL4NP5130
intestinal stem cell of posterior adult midgut epithelium | somatic clone
intestinal stem cell of posterior adult midgut epithelium | somatic clone, with Scer\GAL4Act5C.PP
NMJ bouton | larval stage, with Sdc48/Sdc10608
posterior crossvein
posterior wing margin, with Scer\GAL4vg.int2.1
triple row of wing sensilla, with Scer\GAL4vg.int2.1
wing disc
wing disc (with Df(2R)PK1)
wing vein
wing vein | ectopic
wing vein L5, with Scer\GAL4vg.int2.1
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) (3)
Hsap\ZFYVE9
11 of 14
Yes
Yes
Hsap\ZFYVE16
10 of 14
No
Yes
Hsap\ZFYVE28
1 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) (3)
Rnor\Zfyve9
11 of 14
Yes
Yes
Rnor\Zfyve16
10 of 14
No
Yes
Rnor\Zfyve28
1 of 14
No
No
Mus musculus (laboratory mouse) (4)
Mmus\Zfyve9
11 of 14
Yes
Yes
Mmus\Zfyve16
10 of 14
No
Yes
Mmus\Mtmr3
1 of 14
No
No
Mmus\Zfyve28
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (3)
Xtro\zfyve9
8 of 13
Yes
Yes
Xtro\zfyve16
8 of 13
Yes
Yes
Xtro\actrt2
1 of 13
No
No
Danio rerio (Zebrafish) (4)
Drer\zfyve16
11 of 14
Yes
Yes
Drer\zfyve9a
9 of 14
No
Yes
Drer\zfyve9b
3 of 14
No
Yes
Drer\si:ch211-11n16.2
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (2)
Cele\aka-1
8 of 14
Yes
Yes
Cele\tag-77
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (1)
Agam\AgaP_AGAP004049
11 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (2)
Atha\AT1G29800
1 of 13
Yes
No
Atha\AT3G43230
1 of 13
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (1)
Scer\PIB2
1 of 13
Yes
No
Schizosaccharomyces pombe (Fission yeast) (1)
Spom\SPBC9B6.03
1 of 12
Yes
Yes
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Sara. 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) (3)
CG3632
1 of 13
CG6051
1 of 13
rush
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 ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    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.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    ZFYVE9; zinc finger FYVE-type containing 9
    11 of 14
    603755
        ZFYVE16; zinc finger FYVE-type containing 16
        10 of 14
        608880
            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
            Sara|Sdc
            enhanceable
            Lar
            (Johnson et al., 2006)
            Sara
            suppressible
            N
            (Montagne and Gonzalez-Gaitan, 2014)
            Starting gene(s)
            Interaction type
            Interacting gene(s)
            Reference
            External Data
            Linkouts
            BioGRID - A database of protein and genetic interactions.
            DroID - A comprehensive database of gene and protein interactions.
            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)
            2R
            Recombination map
            2-95
            Cytogenetic map
            57E6-57E6
            Sequence location
            FlyBase Computed Cytological Location
            Cytogenetic map
            Evidence for location
            57E6-57E6
            Limits computationally determined from genome sequence between P{PZ}tmx1&P{EP}MESK2EP2347 and P{PZ}Egfr05351
            Experimentally Determined Cytological Location
            Cytogenetic map
            Notes
            References
            57E1-57E1
            (Martin, 2000.3.1)
            57E1-57E2
            (Martin et al., 1999)
            Experimentally Determined Recombination Data
            Location

            2-95

            (FlyBase, 2016)

            2-101.5

            (Comeron et al., 2012)
            Left of (cM)
            Right of (cM)
            Notes
            Stocks and Reagents
            Stocks (8)
            Genomic Clones (23)
            cDNA Clones (37)
             

            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
             

            polyclonal

            (Coumailleau et al., 2009)
            Commercially Available Antibodies
             
            Cell Line Information
            Publicly Available Cell Lines
             
              Other Stable Cell Lines
               
                Other Comments
                Relationship to Other Genes
                Source for database merge of

                Source for merge of: Sara CG15667

                (FlyBase, 1996-)
                Additional comments

                Source for merge of Sara CG15667 was sequence comparison ( date:000423 ).

                (FlyBase, 1996-)
                Nomenclature History
                Source for database identify of
                Nomenclature comments
                Etymology
                Synonyms and Secondary IDs (7)
                Reported As
                Symbol Synonym
                CG15667
                (Trivedi et al., 2020, Meadows, 2017.1.11, Ni et al., 2011.7.19, Grieder et al., 2007, Bennett et al., 2006, Molnar et al., 2006, Brody et al., 2002)
                SARA
                (Balakrishnan et al., 2015, van de Hoef et al., 2013, Murray et al., 2012, Peterson et al., 2012, Muñoz-Descalzo et al., 2010, Ashe, 2006, Marques, 2005, Gonzalez-Gaitan, 2003, Ashe, 2002)
                Sara
                (Pinot and Le Borgne, 2024, Yee et al., 2019, Liu and Jin, 2017, Nguyen et al., 2016, Derivery et al., 2015, Zhou et al., 2015, Loubéry and González-Gaitán, 2014, Loubéry et al., 2014, Montagne and Gonzalez-Gaitan, 2014, van de Hoef et al., 2013, Couturier et al., 2012, Daskalaki et al., 2011, Halbsgut et al., 2011, Knox et al., 2011, Coumailleau et al., 2009, Zeng et al., 2008, Gonzales-Gaitan, 2007, Grieder et al., 2007, Kirchner, 2007.7.19, Bennett et al., 2006, Bokel et al., 2006, Knoblich, 2006, Jordan et al., 2005, Winkler et al., 2005)
                dSARA
                (Peterson et al., 2012, Chong et al., 2004, Bokel and Gonzalez-Gaitan, 2003, Bennett and Alphey, 2002, Martin et al., 2002, Bennett and Alphey, 2001, Das et al., 2000, Martin et al., 1999)
                sara
                (Issman-Zecharya and Schuldiner, 2014, Knox et al., 2011, Schulz et al., 2011, Bokel et al., 2006, Johnson et al., 2006, Molnar et al., 2006, Lee et al., 2003, Epstein et al., 2002, Epstein et al., 2001)
                Name Synonyms
                Sara
                (Loubéry et al., 2017)
                Smad Anchor for Receptor Activation
                (Peterson et al., 2012)
                Smad anchor for receptor activation
                (Bennett et al., 2006, Jordan et al., 2005, Bennett and Alphey, 2001, Epstein et al., 2001)
                Secondary FlyBase IDs
                • FBgn0034630
                Datasets (0)
                Study focus (0)
                Experimental Role
                Project
                Project Type
                Title
                Study result (0)
                Result
                Result Type
                Title
                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/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
                FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
                iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
                Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
                MIST (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 (88)