FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\Crag
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General Information
Symbol
Dmel\Crag
Species
D. melanogaster
Name
Calmodulin-binding protein related to a Rab3 GDP/GTP exchange protein
Annotation Symbol
CG12737
Feature Type
protein_coding_gene
FlyBase ID
FBgn0025864
Gene Model Status
Current
Stock Availability
18 publicly available
Gene Summary
Calmodulin-binding protein related to a Rab3 GDP/GTP exchange protein (Crag) encodes a guanine exchange factor for the products of Rab10 and Rab11. It is involved in the exclusive secretion of basement membrane proteins on the basal side of cells. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Key Links
Genomic Location
Cytogenetic map
7F7-7F8
Sequence location
Recombination map
1-23
RefSeq locus
NC_004354 REGION:8592031..8602036
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 (17 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
enables calmodulin binding
inferred from direct assay
(Denef et al., 2008)
inferred from physical interaction with FLYBASE:Cam; FB:FBgn0000253
(Xu et al., 1998)
enables guanyl-nucleotide exchange factor activity
inferred from direct assay
(Xiong et al., 2012)
enables small GTPase binding
inferred from physical interaction with FLYBASE:Rab11; FB:FBgn0015790
(Xiong et al., 2012)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
enables guanyl-nucleotide exchange factor activity
inferred from biological aspect of ancestor with PANTHER:PTN000265677
(GO Reference Genome Project, 2011-)
Biological Process (9 terms)
Terms Based on Experimental Evidence (9 terms)
CV Term
Evidence
References
involved_in basal protein localization
inferred from mutant phenotype
(Lerner et al., 2013)
involved_in basement membrane assembly
inferred from mutant phenotype
(Denef et al., 2008)
involved_in basement membrane constituent secretion
inferred from mutant phenotype
(Isabella and Horne-Badovinac, 2016)
involved_in basolateral protein secretion
inferred from mutant phenotype
(Isabella and Horne-Badovinac, 2016)
involved_in defense response to Gram-negative bacterium
inferred from high throughput mutant phenotype
(Cronin et al., 2009)
involved_in positive regulation of Golgi to plasma membrane protein transport
inferred from mutant phenotype
(Xiong et al., 2012)
involved_in positive regulation of innate immune response
inferred from high throughput mutant phenotype
(Cronin et al., 2009)
involved_in regulation of basement membrane organization
inferred from mutant phenotype
(Isabella and Horne-Badovinac, 2016)
involved_in regulation of Rab protein signal transduction
inferred from direct assay
(Xiong et al., 2012)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
involved_in regulation of Rab protein signal transduction
inferred from biological aspect of ancestor with PANTHER:PTN000265677
(GO Reference Genome Project, 2011-)
Cellular Component (5 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
located_in apical cortex
inferred from direct assay
(Devergne et al., 2017, Denef et al., 2008)
located_in cytoplasmic vesicle
inferred from direct assay
(Xiong et al., 2012)
located_in lateral cell cortex
inferred from direct assay
(Devergne et al., 2017, Denef et al., 2008)
located_in plasma membrane
inferred from direct assay
(Devergne et al., 2014)
located_in recycling endosome
inferred from direct assay
(Denef et al., 2008)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in cytoplasmic vesicle
inferred from biological aspect of ancestor with PANTHER:PTN000265677
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
Calmodulin-binding protein related to a Rab3 GDP/GTP exchange protein (Crag) encodes a guanine exchange factor for the products of Rab10 and Rab11. It is involved in the exclusive secretion of basement membrane proteins on the basal side of cells. [Date last reviewed: 2019-03-07]
Gene Group (FlyBase)
RAB GUANINE NUCLEOTIDE EXCHANGE FACTORS - DENN DOMAIN -
DENN domain-containing proteins are Guanine Nucleotide Exchange Factors (GEFs) for Rab family GTPases. (Adapted from FBrf0220596).
Protein Function (UniProtKB)
Calmodulin-binding protein that acts as a guanine exchange factor for Rab10 and Rab11 (PubMed:23226104, PubMed:23369713, PubMed:9813038). Essential for maintenance of adult photoreceptor cells (PubMed:23226104). Upon light stimulation, required for trafficking of newly synthesized ninaE (Rh1) from the trans-Golgi network to rhabdomere membranes via Rab11-dependent vesicular transport (PubMed:23226104). During egg development, essential for establishing and maintaining epithelial cell polarity by regulating the correct polarized deposition of basal membrane (BM) proteins in follicular epithelial (FE) cells (PubMed:18331716, PubMed:23369713, PubMed:24828534). Functions by targeting Rab10 to the basal cytoplasm, where it restricts the secretion of BM proteins such as trol/Pcan and vkg/Coll IV to the basal surface (PubMed:18331716, PubMed:23369713, PubMed:24828534). Appears to be involved in regulating the levels and distribution of the guanine nucleotide exchange factor strat, however the two proteins appear to have independent roles in regulating polarized BM protein secretion in the FE (PubMed:28228250).
(UniProt, Q9W3D3)
Summary (Interactive Fly)

guanine exchange factor for Rab10 and Rab11 that organizes epithelial architecture by regulating the polarized secretion of basement membrane proteins

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

Please see the JBrowse view of Dmel\Crag 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 Q9W3D3)

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 5.52

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Crag-RA
FBtr0071219
NM_080341
6196
1671
Crag-RB
FBtr0071220
NM_167164
6115
1644
Crag-RD
FBtr0343356
NM_001169239
7001
1671
Additional Transcript Data and Comments
Reported size (kB)

4.6, 2.5 (northern blot)

(Xu et al., 1998)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
Crag-PA
FBpp0071165
187.0
1671
6.36
Q9W3D3
NP_525080
AAF46398
Crag-PB
FBpp0089320
184.0
1644
6.28
Q9W3D2
NP_727281
AAF46397
Crag-PD
FBpp0310013
187.0
1671
6.36
Q9W3D3
NP_001162710
ACZ95245
Polypeptides with Identical Sequences

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

1671 aa isoforms: Crag-PA, Crag-PD
Additional Polypeptide Data and Comments
Reported size (kDa)

1441 (aa)

(Xu et al., 1998)
Comments
External Data
Subunit Structure (UniProtKB)

Interacts with Cam (PubMed:18331716, PubMed:9813038). Interacts with Rab10 (PubMed:23369713). Interacts (via the DENN domains) with Rab11 (PubMed:23226104).

(UniProt, Q9W3D3)
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\Crag using the Feature Mapper tool.

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

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

-1.22

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
organism
(Fisher et al., 2012)
embryonic stage 7 -- 8
organism | ubiquitous
(Fisher et al., 2012)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
(Xu et al., 1998)
Additional Descriptive Data

Crag transcripts are expressed at high levels in adult flies,and at very low levels in other developmental stages.

(Xu et al., 1998)

Crag transcripts are expressed at high levels in adult flies, and at very low levels in other developmental stages.

(Xu et al., 1998)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
oogenesis
germarium

Comment: in lateral and apical cortex

(Denef et al., 2008)
follicle cell

Comment: in lateral and apical cortex

(Denef et al., 2008)
follicle cell

Comment: !

(Denef et al., 2008)
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult head
(Aradska et al., 2015)
Additional Descriptive Data

Crag protein is observed in the follicular epithelium as it is formed in the germarium and continues to be expressed in the entire epithelium throughout its development. Crag protein accumulates at the plasma membrane and in punctate structures throughout the cytoplasm, which were identified as endosomes. In early stages it is present along the lateral and apical cortex and later is more prominent in the lateral cortex. It is not observed in the basal cortex.

(Denef et al., 2008)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in apical cortex
inferred from direct assay
(Devergne et al., 2017, Denef et al., 2008)
located_in cytoplasmic vesicle
inferred from direct assay
(Xiong et al., 2012)
located_in lateral cell cortex
inferred from direct assay
(Devergne et al., 2017, Denef et al., 2008)
located_in plasma membrane
inferred from direct assay
(Devergne et al., 2014)
located_in recycling endosome
inferred from direct assay
(Denef et al., 2008)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Crag 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
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
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 12 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 9 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Crag
Transgenic constructs containing regulatory region of Crag
Aberrations (Deficiencies and Duplications) ( 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 - all die before end of larval stage (with CragA)
lethal - all die before end of larval stage (with CragC)
lethal - all die before end of larval stage (with CragD)
lethal - all die before end of larval stage (with Df(1)BSC627)
lethal - all die before end of larval stage (with Df(1)KA14)
lethal - all die before end of larval stage | recessive
lethal - all die before end of second instar larval stage (with CragA)
lethal - all die before end of second instar larval stage (with CragC)
lethal - all die before end of second instar larval stage (with Df(1)BSC627)
lethal - all die before end of second instar larval stage | recessive
partially lethal - majority die, with Scer\GAL4pnr-MD237
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4tin.CΔ4
Other Phenotypes
Allele
abnormal body color, with Scer\GAL4pnr-MD237
abnormal neuroanatomy | larval stage
abnormal neurophysiology | adult stage | progressive | somatic clone | conditional
abnormal neurophysiology | adult stage | somatic clone
abnormal wound healing | larval stage
short lived | heat sensitive, with Scer\GAL4tin.CΔ4
some die during larval stage (with Df(1)KA14)
some die during larval stage | recessive
some die during pupal stage, with Scer\GAL4pnr-MD237
some die during second instar larval stage (with CragA)
some die during second instar larval stage (with CragC)
some die during second instar larval stage (with Df(1)BSC627)
some die during second instar larval stage | recessive
some die during third instar larval stage (with CragA)
some die during third instar larval stage (with CragC)
some die during third instar larval stage (with CragD)
some die during third instar larval stage (with Df(1)BSC627)
some die during third instar larval stage | recessive
visible, with Scer\GAL4pnr-MD237
Phenotype manifest in
Allele
axon | larval stage
follicle cell, with Scer\GAL4tj.PU
follicle cell | somatic clone
larval multidendritic class III neuron
mesonotum, with Scer\GAL4pnr-MD237
ovarian basement membrane | oogenesis, with Scer\GAL4tj.PU
rhabdomere | progressive | somatic clone | conditional
trans-Golgi network | somatic clone
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) (15)
Hsap\DENND4A
13 of 14
Yes
Yes
 
1  
Hsap\DENND4B
13 of 14
Yes
Yes
Hsap\DENND4C
13 of 14
Yes
Yes
Hsap\DENND3
4 of 14
No
Yes
Hsap\SBF1
2 of 14
No
No
Hsap\DENND1A
1 of 14
No
No
1  
Hsap\DENND1B
1 of 14
No
No
Hsap\DENND1C
1 of 14
No
No
Hsap\DENND2A
1 of 14
No
No
Hsap\DENND2B
1 of 14
No
No
Hsap\DENND2C
1 of 14
No
No
Hsap\DENND2D
1 of 14
No
No
Hsap\DENND5A
1 of 14
No
No
Hsap\DENND5B
1 of 14
No
No
Hsap\SBF2
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) (13)
Rnor\Dennd4a
13 of 14
Yes
Yes
Rnor\Dennd4b
13 of 14
Yes
Yes
Rnor\Dennd4c
12 of 14
No
Yes
Rnor\Dennd3
4 of 14
No
Yes
Rnor\Sbf1
2 of 14
No
No
Rnor\Dennd1a
1 of 14
No
No
Rnor\Dennd1b
1 of 14
No
No
Rnor\Dennd1c
1 of 14
No
No
Rnor\Dennd2a
1 of 14
No
No
Rnor\Dennd2b
1 of 14
No
Yes
Rnor\Dennd2c
1 of 14
No
No
Rnor\Dennd2d
1 of 14
No
No
Rnor\Dennd5a
1 of 14
No
No
Mus musculus (laboratory mouse) (17)
Mmus\Dennd4a
13 of 14
Yes
Yes
Mmus\Dennd4b
13 of 14
Yes
Yes
Mmus\Dennd4c
13 of 14
Yes
Yes
Mmus\Dennd3
4 of 14
No
Yes
Mmus\Sbf1
2 of 14
No
No
Mmus\Apaf1
1 of 14
No
No
Mmus\Denn2b
1 of 14
No
No
Mmus\Dennd1a
1 of 14
No
No
Mmus\Dennd1b
1 of 14
No
No
Mmus\Dennd1c
1 of 14
No
No
Mmus\Dennd2a
1 of 14
No
No
Mmus\Dennd2c
1 of 14
No
No
Mmus\Dennd2d
1 of 14
No
No
Mmus\Dennd5a
1 of 14
No
No
Mmus\Dennd5b
1 of 14
No
No
Mmus\Madd
1 of 14
No
No
Mmus\Sbf2
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (9)
Xtro\dennd4c
9 of 13
Yes
Yes
Xtro\dennd4a
6 of 13
No
Yes
Xtro\dennd4b
5 of 13
No
Yes
Xtro\ranbp9
5 of 13
No
No
Xtro\dennd2c
1 of 13
No
Yes
Xtro\madd
1 of 13
No
No
Xtro\mapre2
1 of 13
No
No
Xtro\mreg
1 of 13
No
Yes
Xtro\sbf1
1 of 13
No
No
Danio rerio (Zebrafish) (15)
Drer\dennd4a
12 of 14
Yes
Yes
Drer\dennd4c
12 of 14
Yes
Yes
Drer\dennd4b
10 of 14
No
Yes
Drer\dennd3a
4 of 14
No
Yes
Drer\dennd1a
1 of 14
No
No
Drer\dennd1b
1 of 14
No
No
Drer\dennd1c
1 of 14
No
No
Drer\dennd2b
1 of 14
No
Yes
Drer\dennd2c
1 of 14
No
No
Drer\dennd2da
1 of 14
No
Yes
Drer\dennd2db
1 of 14
No
Yes
Drer\dennd5a
1 of 14
No
No
Drer\dennd5b
1 of 14
No
No
Drer\sbf1
1 of 14
No
No
Drer\sbf2
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (5)
Cele\denn-4
12 of 14
Yes
Yes
Cele\lst-6
2 of 14
No
No
Cele\aex-3
1 of 14
No
No
Cele\mtm-5
1 of 14
No
No
Cele\rme-4
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (4)
Agam\AgaP_AGAP001102
12 of 12
Yes
Yes
Agam\AgaP_AGAP005212
1 of 12
No
No
Agam\AgaP_AGAP009774
1 of 12
No
No
Agam\AgaP_AGAP011351
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (2)
Atha\SCD1
5 of 13
Yes
Yes
Atha\AT5G35560
1 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (2)
Spom\SPCC297.05
2 of 12
Yes
Yes
Spom\SPBC3D6.12
1 of 12
No
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Crag. 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)
CG18659
1 of 13
pns
1 of 13
Sbf
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?
    DENND4A; DENN domain containing 4A
    13 of 14
    600382
         
        DENND4B; DENN domain containing 4B
        13 of 14
        619843
            DENND4C; DENN domain containing 4C
            13 of 14
                DENND3; DENN domain containing 3
                4 of 14
                617503
                    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.
                    Dmel gene
                    Ortholog showing functional complementation
                    Supporting References
                    Crag
                    Hsap\DENND4A
                    (Xiong et al., 2012)
                    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
                    Crag
                    suppressible
                    Rab11
                    (Xiong et al., 2012)
                    Starting gene(s)
                    Interaction type
                    Interacting gene(s)
                    Reference
                    External Data
                    Subunit Structure (UniProtKB)
                    Interacts with Cam (PubMed:18331716, PubMed:9813038). Interacts with Rab10 (PubMed:23369713). Interacts (via the DENN domains) with Rab11 (PubMed:23226104).
                    (UniProt, Q9W3D3 )
                    Linkouts
                    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
                    Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                    Class of Gene
                    Genomic Location and Detailed Mapping Data
                    Chromosome (arm)
                    X
                    Recombination map
                    1-23
                    Cytogenetic map
                    7F7-7F8
                    Sequence location
                    FlyBase Computed Cytological Location
                    Cytogenetic map
                    Evidence for location
                    7F7-7F8
                    Limits computationally determined from genome sequence between P{EP}CG1632EP1583 and P{EP}MoeEP1652
                    Experimentally Determined Cytological Location
                    Cytogenetic map
                    Notes
                    References
                    8A1-8A2
                    (Montell, 1998.8.11)
                    8A1-8A2
                    (determined by in situ hybridisation)
                    (Xu et al., 1998)
                    Experimentally Determined Recombination Data
                    Location

                    1-23

                    (FlyBase, 2016)

                    1-23.7

                    (Comeron et al., 2012)
                    Left of (cM)
                    Right of (cM)
                    Notes
                    Stocks and Reagents
                    Stocks (18)
                    Genomic Clones (10)
                     

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

                    cDNA Clones (98)
                     

                    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

                      (Devergne et al., 2017, Denef et al., 2008)
                      Commercially Available Antibodies
                       
                      Cell Line Information
                      Publicly Available Cell Lines
                       
                        Other Stable Cell Lines
                         
                          Other Comments

                          Crag is a guanyl-nucleotide exchange factor for Rab11 and is required for the post-Golgi transport of ninaE to the rhabdomeres during light activation.

                          (Xiong et al., 2012)

                          Crag plays a role in organising epithelial architecture by regulating polarised secretion of basement membrane proteins.

                          (Denef et al., 2008)

                          Identification: Cam-binding protein that is expressed in the retina.

                          (Xu et al., 1998)
                          Relationship to Other Genes
                          Source for database merge of
                          Additional comments
                          Nomenclature History
                          Source for database identify of
                          Nomenclature comments
                          Etymology
                          Synonyms and Secondary IDs (7)
                          Datasets (0)
                          Study focus (0)
                          Experimental Role
                          Project
                          Project Type
                          Title
                          Study result (0)
                          Result
                          Result Type
                          Title
                          External Crossreferences and Linkouts ( 70 )
                          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
                          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
                          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
                          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
                          MIST (protein-protein) - An integrated Molecular Interaction Database
                          Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                          References (65)