FB2026_02 , released June 18, 2026
Gene: Dmel\fra
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General Information
Symbol
Dmel\fra
Species
D. melanogaster
Name
frazzled
Annotation Symbol
CG8581
Feature Type
protein_coding_gene
FlyBase ID
FBgn0011592
Gene Model Status
Current
Stock Availability
71 publicly available
Gene Summary
frazzled (fra) encodes a DCC-like Netrin receptor that mediates axon guidance. It also contributes to dendrite guidance, development and morphogenesis. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Also Known As

DCC

Key Links
Genomic Location
Cytogenetic map
49A10-49B3
Sequence location
Recombination map
2-67
RefSeq locus
NT_033778 REGION:12529053..12564184
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 (23 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables protein domain specific binding
inferred from physical interaction with UniProtKB:M9MRI4
(Morikawa et al., 2011)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
enables netrin receptor activity
traceable author statement
(Bashaw and Goodman, 1999)
Biological Process (14 terms)
Terms Based on Experimental Evidence (12 terms)
CV Term
Evidence
References
involved_in axon guidance
inferred from mutant phenotype
(O'Donnell and Bashaw, 2013, Yang et al., 2009, Garbe et al., 2007, Furrer et al., 2003)
inferred from genetic interaction with FLYBASE:comm; FB:FBgn0010105
(Yang et al., 2009)
inferred from genetic interaction with FLYBASE:robo1; FB:FBgn0005631
(Hiramoto and Hiromi, 2006)
involved_in axon midline choice point recognition
inferred from genetic interaction with UniProtKB:M9MRI4
(Song et al., 2011)
inferred from mutant phenotype
(Morikawa et al., 2011)
inferred from mutant phenotype
(Ratnaparkhi et al., 2002)
involved_in compound eye development
inferred from mutant phenotype
(Akin and Zipursky, 2016)
inferred from genetic interaction with UniProtKB:Q24567,UniProtKB:Q24568
(Akin and Zipursky, 2016)
involved_in dendrite guidance
inferred from mutant phenotype
(Furrer et al., 2003)
involved_in dendrite morphogenesis
inferred from mutant phenotype
(Mauss et al., 2009)
involved_in glial cell migration
inferred from mutant phenotype
(von Hilchen et al., 2010)
involved_in motor neuron axon guidance
inferred from mutant phenotype
(Dorsten et al., 2007, Labrador et al., 2005, Kolodziej et al., 1996)
involved_in peripheral nervous system neuron axonogenesis
inferred from mutant phenotype
(Morikawa et al., 2011)
inferred from genetic interaction with UniProtKB:M9MRI4
(Morikawa et al., 2011)
involved_in positive regulation of cell migration
inferred from mutant phenotype
(Raza and Jacobs, 2016)
involved_in regulation of axonogenesis
inferred from mutant phenotype
(Morikawa et al., 2011)
inferred from genetic interaction with UniProtKB:M9MRI4
(Morikawa et al., 2011)
involved_in regulation of dendrite development
inferred from mutant phenotype
(Brierley et al., 2009)
involved_in salivary gland boundary specification
inferred from mutant phenotype
(Kolesnikov and Beckendorf, 2005)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
involved_in axon guidance
traceable author statement
(Grueber and Jan, 2004)
inferred from biological aspect of ancestor with PANTHER:PTN001543908
(GO Reference Genome Project, 2011-)
involved_in cell-cell adhesion
inferred from biological aspect of ancestor with PANTHER:PTN002548291
(GO Reference Genome Project, 2011-)
involved_in dendrite morphogenesis
traceable author statement
(Grueber and Jan, 2004)
involved_in neuron migration
inferred from biological aspect of ancestor with PANTHER:PTN001543908
(GO Reference Genome Project, 2011-)
Cellular Component (7 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
located_in axonal growth cone
inferred from mutant phenotype
(Akin and Zipursky, 2016)
inferred from genetic interaction with UniProtKB:Q24567,UniProtKB:Q24568
(Akin and Zipursky, 2016)
located_in axon
inferred from direct assay
(Furrer et al., 2003)
located_in dendrite
inferred from direct assay
(Furrer et al., 2003)
located_in neuronal cell body
inferred from direct assay
(Furrer et al., 2003)
located_in perivitelline space
inferred from high throughput direct assay
(Lye et al., 2014)
located_in plasma membrane
inferred from high throughput direct assay
(Tan et al., 2009)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
is_active_in axonal growth cone
inferred from biological aspect of ancestor with PANTHER:PTN001543908
(GO Reference Genome Project, 2011-)
located_in membrane
inferred from electronic annotation with InterPro:IPR010560
(InterPro Project Members, 2004-)
is_active_in plasma membrane
inferred from biological aspect of ancestor with PANTHER:PTN001543908
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
frazzled (fra) encodes a DCC-like Netrin receptor that mediates axon guidance. It also contributes to dendrite guidance, development and morphogenesis. [Date last reviewed: 2019-03-07]
Gene Group (FlyBase)
UNCLASSIFIED TRANSMEMBRANE RECEPTORS -
This group comprises transmembrane receptors that do not classify under other groups in FlyBase.
Summary (Interactive Fly)

Ig superfamily - functions as a Netrin receptor in axon pathfinding - promotes growth cone attachment at the source of a Netrin gradient in the visual system

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

Please see the JBrowse view of Dmel\fra 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 A1Z920)

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 5.49

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
fra-RA
FBtr0087822
NM_078992
6320
1526
fra-RB
FBtr0087821
NM_165913
5867
1375
fra-RC
FBtr0339649
NM_001299424
6293
1517
Additional Transcript Data and Comments
Reported size (kB)

6.318, 5.865 (compiled cDNA)

(Kolodziej et al., 1996)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
fra-PA
FBpp0086935
168.8
1526
6.49
A1Z920
NP_523716
AAF58493
fra-PB
FBpp0086934
151.7
1375
6.54
A1Z921
NP_725185
AAM68622
fra-PC
FBpp0308711
167.7
1517
6.25
A0A0B4LEX6
NP_001286353
AHN56151
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)

1526, 1375 (aa)

(Kolodziej et al., 1996)
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\fra 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).

-0.43

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
presumptive embryonic/larval central nervous system
(Kolodziej et al., 1996)
larval digestive system
(Kolodziej et al., 1996)
epithelial cell

Comment: clustered apical distribution

(Lecuyer et al., 2007)
epithelium
(Lecuyer et al., 2007)
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 7 -- 8
antennal anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
central brain anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
dorsal head epidermis anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
visual anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
dorsal ectoderm
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 9 -- 10
antennal primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
central brain primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
visual primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
dorsal head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
lateral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
ventral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
dorsal ectoderm
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 10
lateral ectoderm
(von Hilchen et al., 2010)
embryonic stage 11
longitudinal glial cell
(von Hilchen et al., 2010)
ventral ectoderm | segmentally repeated
(Joly et al., 2007)
embryonic stage 11 -- 12
anterior midgut primordium
(Fisher et al., 2012)
central brain primordium
(Fisher et al., 2012)
dorsal epidermis primordium
(Fisher et al., 2012)
dorsal head epidermis primordium

Comment: reported as head epidermis primordium

(Fisher et al., 2012)
lateral head epidermis primordium

Comment: reported as head epidermis primordium

(Fisher et al., 2012)
ventral head epidermis primordium

Comment: reported as head epidermis primordium

(Fisher et al., 2012)
embryonic optic lobe primordium
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
posterior midgut primordium
(Fisher et al., 2012)
ventral epidermis primordium
(Fisher et al., 2012)
ventral nerve cord primordium
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic brain
(Fisher et al., 2012)
embryonic dorsal epidermis
(Fisher et al., 2012)
embryonic head epidermis
(Fisher et al., 2012)
embryonic proventriculus inner layer
(Fisher et al., 2012)
embryonic ventral epidermis
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
larval nervous system
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
sensory nervous system primordium
(Fisher et al., 2012)
larval ventral nerve cord
(Fisher et al., 2012)
Additional Descriptive Data

fra transcripts are abundantly expressed in the CNS and the gut epithelium.

(Kolodziej et al., 1996)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 10
lateral ectoderm
(von Hilchen et al., 2010)
embryonic stage 12 -- 17
embryonic/larval midgut
(Kolodziej et al., 1996)
embryonic stage 13 -- 17
larval ventral nerve cord commissure
(Kolodziej et al., 1996)
neuron
• axon
(Kolodziej et al., 1996)
embryonic stage 15
larval segmental nerve
(Kolodziej et al., 1996)
larval longitudinal connective
(Kolodziej et al., 1996)
motor neuron
(Kolodziej et al., 1996)
larval intersegmental nerve
(Kolodziej et al., 1996)
epidermis
(Kolodziej et al., 1996)
third instar larval stage
embryonic/larval optic stalk
(Joly et al., 2007)
lamina anlage
(Joly et al., 2007)
wing disc
(Joly et al., 2007)
pupal stage P6
photoreceptor cell R8

Comment: 24h APF

(Timofeev et al., 2012)
pupal stage P6 -- P7
medulla anlage | distal | restricted

Comment: 24-42h APF

(Timofeev et al., 2012)
pupal stage P6 -- pharate adult stage P8
retina | restricted

Comment: 24-55h APF

(Timofeev et al., 2012)
photoreceptor cell R8
• axon

Comment: 24-55h APF

(Timofeev et al., 2012)
pupal stage P7
rhabdomere R1

Comment: 42h APF

(Timofeev et al., 2012)
rhabdomere R2

Comment: 42h APF

(Timofeev et al., 2012)
rhabdomere R3

Comment: 42h APF

(Timofeev et al., 2012)
rhabdomere R4

Comment: 42h APF

(Timofeev et al., 2012)
rhabdomere R5

Comment: 42h APF

(Timofeev et al., 2012)
rhabdomere R6

Comment: 42h APF

(Timofeev et al., 2012)
pupal stage P7 -- pharate adult stage P8
rhabdomere R8

Comment: 42-55h APF

(Timofeev et al., 2012)
pharate adult stage P8
medulla layer M3

Comment: 55h APF

(Timofeev et al., 2012)
adult stage
medulla layer M3 | faint
(Timofeev et al., 2012)
adult optic lobe glial cell of medulla
(Timofeev et al., 2012)
adult optic lobe glial cell of lamina
(Timofeev et al., 2012)
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult head
(Aradska et al., 2015)
Additional Descriptive Data

fra protein is expressed in the pupal retina and medulla. In the retina, the protein is detected in the cell body of photoreceptor R8 cells at 24h APF. After that it is detected in the R8 rhadbomere. Transient expression in rhabdomeres R1-R6 is also seen at 42h APF. In the medulla, fra accumulates at the distal border at 24-42h APF, where R8 axons temporarily pause. After that it localises to medulla layer M3, the layer targeted by R8 axons. Weak expression persists in adults. Expression of fra is also detected in glial cell subtypes in the adult medulla and lamina.

(Timofeev et al., 2012)

fra protein is expressed on developing axons and epithelia in the embyro. It is detected at high levels on commissural and longitudinal axons and at lower levels on peripheral motor axons that extend outward in the segmental and intersegmental nerves. It is also detected on the surfaces of midgut epithelial cells and on epidermis.

(Kolodziej et al., 1996)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in axonal growth cone
inferred from mutant phenotype
(Akin and Zipursky, 2016)
inferred from genetic interaction with UniProtKB:Q24567,UniProtKB:Q24568
(Akin and Zipursky, 2016)
located_in axon
inferred from direct assay
(Furrer et al., 2003)
located_in dendrite
inferred from direct assay
(Furrer et al., 2003)
located_in neuronal cell body
inferred from direct assay
(Furrer et al., 2003)
located_in perivitelline space
inferred from high throughput direct assay
(Lye et al., 2014)
located_in plasma membrane
inferred from high throughput direct assay
(Tan et al., 2009)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\fra 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
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 9-10
  • Stages(s) 11-12
  • Stages(s) 13-16
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 20 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 49 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of fra
Transgenic constructs containing regulatory region of fra
Aberrations (Deficiencies and Duplications) ( 3 )
Inferred from experimentation ( 3 )
Gene disrupted in
Df(2R)vg135
(Hummel et al., 1999)
Df(2R)vg-C
(Ratnaparkhi et al., 2002)
T(2;3)fra1
(Kolodziej et al., 1996)
Inferred from location ( 3 )
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 during embryonic stage | recessive
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4tin.CΔ4
Sterility
Allele
female fertile, with Scer\GAL4nanos.PG
female sterile, with Scer\GAL4nanos.PG
Other Phenotypes
Allele
abnormal behavior
abnormal cell adhesion | embryonic stage
abnormal cell migration | dominant | embryonic stage
abnormal cell migration | third instar larval stage (with fra3)
abnormal cell migration | third instar larval stage (with fra4)
abnormal flight, with Scer\GAL4Mef2.PR
abnormal neuroanatomy
abnormal neuroanatomy, with Scer\GAL4ap-md544
abnormal neuroanatomy, with Scer\GAL4eg-Mz360
abnormal neuroanatomy, with Scer\GAL4eve.CQ2
abnormal neuroanatomy (with Df(2R)BSC3)
abnormal neuroanatomy (with Df(2R)vg135)
abnormal neuroanatomy (with fra3)
abnormal neuroanatomy (with fra4)
abnormal neuroanatomy (with fra6)
abnormal neuroanatomy | adult stage, with Scer\GAL4Pdf.PU
abnormal neuroanatomy | embryonic stage
abnormal neuroanatomy | embryonic stage, with Scer\GAL4ap.PU
abnormal neuroanatomy | embryonic stage, with Scer\GAL4ap-md544
abnormal neuroanatomy | embryonic stage, with Scer\GAL4eg.PU
abnormal neuroanatomy | embryonic stage, with Scer\GAL4eg-Mz360
abnormal neuroanatomy | embryonic stage, with Scer\GAL4eg-Mz360, fraunspecified
abnormal neuroanatomy | embryonic stage, with Scer\GAL4eg-Mz360, fraΔC.UAS.Tag:HA
abnormal neuroanatomy | embryonic stage, with Scer\GAL4elav.PLu
abnormal neuroanatomy | embryonic stage, with Scer\GAL4elav.PLu, fraunspecified
abnormal neuroanatomy | embryonic stage, with Scer\GAL4elav.PLu, fraΔC.UAS.Tag:HA
abnormal neuroanatomy | embryonic stage (with fra3)
abnormal neuroanatomy | embryonic stage (with fra4)
abnormal neuroanatomy | embryonic stage (with fra6)
abnormal neuroanatomy | embryonic stage (with fra23H9)
abnormal neuroanatomy | embryonic stage (with fraGA957)
abnormal neuroanatomy | embryonic stage | recessive
abnormal neuroanatomy | somatic clone
abnormal neuroanatomy | somatic clone, with Scer\GAL4VGlut1-OK371
abnormal neuroanatomy | third instar larval stage (with fra3)
abnormal neuroanatomy | third instar larval stage (with fra4)
abnormal size | adult stage, with Scer\GAL4Pdf.PU
increased cell number | somatic clone
increased cell number | somatic clone, with Scer\GAL4wor.PA
some die during embryonic stage | recessive
Phenotype manifest in
Allele
abdominal 1 ventral monoscolopidial chordotonal organ vch1 | third instar larval stage (with fra3)
abdominal 1 ventral monoscolopidial chordotonal organ vch1 | third instar larval stage (with fra4)
abdominal lch1 neuron | third instar larval stage (with fra3)
abdominal lch1 neuron | third instar larval stage (with fra4)
abdominal ventral monoscolopidial chordotonal organ vch2 | third instar larval stage (with fra3)
abdominal ventral monoscolopidial chordotonal organ vch2 | third instar larval stage (with fra4)
adult optic lobe | somatic clone
adult optic lobe | somatic clone, with Scer\GAL80ey.3.5
adult s-LNv neuron | adult stage, with Scer\GAL4Pdf.PU
axon | adult stage, with Scer\GAL4Pdf.PU
axon | embryonic stage, with Scer\GAL4ap-md544
axon | embryonic stage, with Scer\GAL4eg-Mz360
axon | embryonic stage (with fra3)
axon | embryonic stage (with fra4)
axon | embryonic stage (with fra6)
Bolwig nerve (with fra3)
Bolwig nerve (with fra4)
commissure & embryo & axon
commissure | embryonic stage
commissure | embryonic stage, with Scer\GAL4eg.PU
commissure | embryonic stage (with fra3)
commissure | embryonic stage (with fra4)
commissure | embryonic stage (with fra6)
connective | embryonic stage
connective | embryonic stage (with fra3)
connective | embryonic stage (with fra4)
connective | embryonic stage (with fra6)
dendrite (with fra3)
dendrite (with fra4)
dendrite | somatic clone
dendrite | somatic clone, with Scer\GAL4VGlut1-OK371
dMP2 neuron, with Scer\GAL4605, fra3/fra4
dMP2 neuron (with fra3)
dMP2 neuron (with fra3), with Scer\GAL4605, fraΔC.UAS.Tag:MYC
dMP2 neuron (with fra4)
dMP2 neuron (with fra4), with Scer\GAL4605, fraΔC.UAS.Tag:MYC
EG neuron, with Scer\GAL4eg-Mz360, fraunspecified
EG neuron, with Scer\GAL4eg-Mz360, fraΔC.UAS.Tag:HA
embryonic heart cardioblast
embryonic heart cardioblast, with Scer\GAL4Mef2.247
embryonic longitudinal glial cell (with fra3)
embryonic longitudinal glial cell (with fra4)
fascicle | embryonic stage, with Scer\GAL4ap.PU
fascicle | embryonic stage (with fra3)
fascicle | embryonic stage (with fra4)
filopodium | embryonic stage
filopodium | embryonic stage, with Scer\GAL4Mef2.247
heart primordium
intermediate neuronal progenitor | increased number | somatic clone
lamellipodium | embryonic stage
lamellipodium | embryonic stage, with Scer\GAL4Mef2.247
lamina
larval anterior commissure
larval anterior commissure (with fra3)
larval anterior commissure (with fra4)
larval anterior commissure (with fra23H9)
larval anterior commissure (with fraGA957)
larval EW neuron
larval EW neuron, with Scer\GAL4ap-md544
larval EW neuron, with Scer\GAL4eg-Mz360
larval EW neuron, with Scer\GAL4eg-Mz360, fraunspecified
larval EW neuron, with Scer\GAL4eg-Mz360, fraΔC.UAS.Tag:HA
larval EW neuron (with Df(2R)BSC3)
larval EW neuron (with Df(2R)vg135)
larval EW neuron (with fra3)
larval EW neuron (with fra4)
larval EW neuron (with fra6)
larval EW neuron | embryonic stage
larval EW neuron | embryonic stage, with Scer\GAL4eg.PU
larval EW neuron | embryonic stage, with Scer\GAL4eg-Mz360
larval EW neuron | embryonic stage (with fra3)
larval EW neuron | embryonic stage (with fra4)
larval EW neuron | embryonic stage (with fra6)
larval interneuron | embryonic stage, with Scer\GAL4ap-md544
larval intersegmental nerve
larval longitudinal connective
larval longitudinal connective (with Df(2R)vg135)
larval longitudinal connective (with fra3)
larval longitudinal connective (with fra4)
larval longitudinal connective (with fra6)
larval longitudinal connective | embryonic stage
larval multidendritic class IV neuron | third instar larval stage (with fra3)
larval multidendritic class IV neuron | third instar larval stage (with fra4)
larval posterior commissure
larval posterior commissure (with Df(2R)vg135)
larval posterior commissure (with fra3)
larval posterior commissure (with fra4)
larval posterior commissure (with fra6)
larval posterior commissure (with fra23H9)
larval posterior commissure (with fraGA957)
larval ventral nerve cord | embryonic stage
larval ventral nerve cord commissure
larval ventral nerve cord commissure (with fra3)
larval ventral nerve cord commissure (with fra4)
larval ventral nerve cord commissure | embryonic stage
larval ventral nerve cord commissure | embryonic stage, with Scer\GAL4elav.PLu
larval ventral nerve cord commissure | embryonic stage, with Scer\GAL4elav.PLu, fraunspecified
larval ventral nerve cord commissure | embryonic stage, with Scer\GAL4elav.PLu, fraΔC.UAS.Tag:HA
larval ventral nerve cord commissure | embryonic stage (with fra3)
larval ventral nerve cord commissure | embryonic stage (with fra4)
larval ventral nerve cord commissure | third instar larval stage (with fra3)
larval ventral nerve cord commissure | third instar larval stage (with fra4)
larval VL3/4 motor neuron
motor neuron, with Scer\GAL4eve.CQ2
motor neuron (with fra3)
motor neuron (with fra4)
motor neuron | somatic clone
motor neuron | somatic clone, with Scer\GAL4VGlut1-OK371
neuroblast | increased number | somatic clone
neuroblast | increased number | somatic clone, with Scer\GAL4wor.PA
neuropil | somatic clone
neuropil | somatic clone, with Scer\GAL4VGlut1-OK371
pCC neuron & axon
photoreceptor cell R8 stalk | somatic clone
photoreceptor cell R8 stalk | somatic clone, with Scer\GAL80ey.3.5
presumptive embryonic/larval central nervous system
presumptive embryonic salivary gland, with Scer\GAL4prd.RG1
presumptive embryonic salivary gland (with fra3)
presumptive embryonic salivary gland (with fra4)
SP1 neuron
symmetrical commissure
symmetrical commissure, with Scer\GAL4eg-Mz360
symmetrical commissure (with Df(2R)BSC3)
symmetrical commissure (with Df(2R)vg135)
symmetrical commissure (with fra3)
symmetrical commissure (with fra4)
symmetrical commissure (with fra6)
symmetrical commissure | embryonic stage (with fra3)
symmetrical commissure | embryonic stage (with fra4)
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) (68)
Hsap\DCC
12 of 14
Yes
Yes
 
5  
Hsap\NEO1
12 of 14
Yes
Yes
Hsap\IGDCC4
3 of 14
No
Yes
Hsap\IGDCC3
2 of 14
No
No
Hsap\PRTG
2 of 14
No
Yes
Hsap\ROBO2
2 of 14
No
No
Hsap\SDK2
2 of 14
No
No
1  
Hsap\ADAMTSL3
1 of 14
No
No
Hsap\BOC
1 of 14
No
No
2  
Hsap\CDON
1 of 14
No
No
Hsap\CHL1
1 of 14
No
No
Hsap\CNTN2
1 of 14
No
No
Hsap\CNTN3
1 of 14
No
No
Hsap\CNTN4
1 of 14
No
No
Hsap\CNTN5
1 of 14
No
No
Hsap\CNTN6
1 of 14
No
No
Hsap\CSF2RB
1 of 14
No
Yes
Hsap\DSCAM
1 of 14
No
No
3  
Hsap\DSCAML1
1 of 14
No
No
2  
Hsap\FGFR2
1 of 14
No
No
Hsap\FLT4
1 of 14
No
No
Hsap\FN1
1 of 14
No
Yes
Hsap\FNDC1
1 of 14
No
Yes
Hsap\FNDC3A
1 of 14
No
No
Hsap\HMCN1
1 of 14
No
No
Hsap\HMCN2
1 of 14
No
No
Hsap\IGSF10
1 of 14
No
No
Hsap\L1CAM
1 of 14
No
No
11  
Hsap\LRIG1
1 of 14
No
No
Hsap\LSAMP
1 of 14
No
No
Hsap\MXRA5
1 of 14
No
No
Hsap\MXRA8
1 of 14
No
Yes
Hsap\MYOT
1 of 14
No
No
Hsap\NCAM1
1 of 14
No
No
4  
Hsap\NEGR1
1 of 14
No
No
1  
Hsap\NFASC
1 of 14
No
No
Hsap\NRCAM
1 of 14
No
No
Hsap\NTM
1 of 14
No
No
Hsap\OPCML
1 of 14
No
No
Hsap\PALLD
1 of 14
No
No
Hsap\PTK7
1 of 14
No
No
1  
Hsap\PTPRA
1 of 14
No
No
1  
Hsap\PTPRB
1 of 14
No
No
Hsap\PTPRC
1 of 14
No
No
Hsap\PTPRD
1 of 14
No
No
Hsap\PTPRE
1 of 14
No
No
Hsap\PTPRF
1 of 14
No
No
2  
Hsap\PTPRG
1 of 14
No
No
Hsap\PTPRK
1 of 14
No
No
Hsap\PTPRM
1 of 14
No
No
Hsap\PTPRQ
1 of 14
No
No
Hsap\PTPRS
1 of 14
No
No
Hsap\PTPRT
1 of 14
No
No
Hsap\PTPRU
1 of 14
No
No
Hsap\PTPRZ1
1 of 14
No
No
Hsap\PXDN
1 of 14
No
No
3  
Hsap\PXDNL
1 of 14
No
No
1  
Hsap\ROBO1
1 of 14
No
No
4  
Hsap\ROBO3
1 of 14
No
No
Hsap\ROS1
1 of 14
No
No
Hsap\SDK1
1 of 14
No
No
Hsap\SORL1
1 of 14
No
No
Hsap\SUSD1
1 of 14
No
Yes
Hsap\TTN
1 of 14
No
No
Hsap\TYRO3
1 of 14
No
Yes
Hsap\UNC5D
1 of 14
No
No
Hsap\USH2A
1 of 14
No
No
Hsap\WFIKKN2
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) (34)
Rnor\Dcc
12 of 14
Yes
Yes
Rnor\Neo1
12 of 14
Yes
Yes
Rnor\Igdcc3
2 of 14
No
Yes
Rnor\Igdcc4
2 of 14
No
No
Rnor\Boc
1 of 14
No
No
Rnor\Bsg
1 of 14
No
No
Rnor\Cdon
1 of 14
No
No
Rnor\Cntn3
1 of 14
No
No
Rnor\Cntn4
1 of 14
No
No
Rnor\Cntn5
1 of 14
No
No
Rnor\Cntn6
1 of 14
No
No
Rnor\Dscaml1
1 of 14
No
No
Rnor\Dscam
1 of 14
No
No
Rnor\Flt4
1 of 14
No
No
Rnor\Fndc1
1 of 14
No
Yes
Rnor\Hmcn1
1 of 14
No
No
Rnor\Igsf10
1 of 14
No
No
Rnor\L1cam
1 of 14
No
No
Rnor\Mypn
1 of 14
No
No
Rnor\Palldl1
1 of 14
No
No
Rnor\Palld
1 of 14
No
No
Rnor\Prtg
1 of 14
No
Yes
Rnor\Ptk7
1 of 14
No
No
Rnor\Ptprd
1 of 14
No
No
Rnor\Ptprf
1 of 14
No
No
Rnor\Ptprm
1 of 14
No
No
Rnor\Ptprq
1 of 14
No
No
Rnor\Ptprs
1 of 14
No
No
Rnor\Pxdn
1 of 14
No
No
Rnor\Robo1
1 of 14
No
No
Rnor\Robo2
1 of 14
No
No
Rnor\Robo4
1 of 14
No
No
Rnor\Sdk2
1 of 14
No
No
Rnor\Unc5d
1 of 14
No
No
Mus musculus (laboratory mouse) (35)
Mmus\Dcc
12 of 14
Yes
Yes
Mmus\Neo1
12 of 14
Yes
Yes
Mmus\Igdcc3
3 of 14
No
Yes
Mmus\Igdcc4
2 of 14
No
Yes
Mmus\Prtg
2 of 14
No
Yes
Mmus\Boc
1 of 14
No
No
Mmus\Cdon
1 of 14
No
No
Mmus\Cntn3
1 of 14
No
No
Mmus\Cntn4
1 of 14
No
No
Mmus\Cntn5
1 of 14
No
No
Mmus\Dscaml1
1 of 14
No
No
Mmus\Dscam
1 of 14
No
No
1  
Mmus\Flt4
1 of 14
No
No
Mmus\Fndc1
1 of 14
No
Yes
Mmus\Hmcn1
1 of 14
No
No
Mmus\Hmcn2
1 of 14
No
No
Mmus\Igsf9
1 of 14
No
No
Mmus\L1cam
1 of 14
No
No
1  
Mmus\Myot
1 of 14
No
Yes
Mmus\Nphs1
1 of 14
No
No
Mmus\Nrcam
1 of 14
No
No
Mmus\Ntm
1 of 14
No
No
Mmus\Ptprd
1 of 14
No
No
Mmus\Ptprf
1 of 14
No
No
Mmus\Ptprm
1 of 14
No
No
0  
Mmus\Ptprq
1 of 14
No
No
Mmus\Ptprs
1 of 14
No
No
Mmus\Pxdn
1 of 14
No
No
Mmus\Robo1
1 of 14
No
No
Mmus\Robo2
1 of 14
No
No
Mmus\Robo4
1 of 14
No
No
Mmus\Sdk1
1 of 14
No
No
Mmus\Susd1
1 of 14
No
Yes
Mmus\Ttn
1 of 14
No
No
Mmus\Unc5d
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (23)
Xtro\dcc
9 of 13
Yes
Yes
Xtro\neo1
7 of 13
No
Yes
Xtro\dscam
1 of 13
No
No
Xtro\fndc3b
1 of 13
No
No
Xtro\hmcn1
1 of 13
No
No
Xtro\hmcn2
1 of 13
No
No
Xtro\igdcc3
1 of 13
No
No
Xtro\iglon5
1 of 13
No
No
Xtro\igsf10
1 of 13
No
No
Xtro\lrig1
1 of 13
No
No
Xtro\musk
1 of 13
No
No
Xtro\ncam2
1 of 13
No
No
Xtro\npffr1.1
1 of 13
No
No
Xtro\nrcam
1 of 13
No
No
Xtro\prtg
1 of 13
No
No
Xtro\ptprt
1 of 13
No
No
Xtro\ptpru
1 of 13
No
No
Xtro\ptprz1
1 of 13
No
No
Xtro\pxdn
1 of 13
No
No
Xtro\sorl1
1 of 13
No
No
Xtro\tyro3
1 of 13
No
Yes
Xtro\ush2a
1 of 13
No
No
Xtro\XB5959262
1 of 13
No
Yes
Danio rerio (Zebrafish) (52)
Drer\LOC100332721
12 of 14
Yes
Yes
Drer\neo1a
12 of 14
Yes
Yes
Drer\dcc
10 of 14
No
Yes
Drer\neo1b
10 of 14
No
Yes
Drer\igdcc3
3 of 14
No
Yes
Drer\igdcc4
3 of 14
No
Yes
Drer\prtgb
3 of 14
No
Yes
Drer\prtga
2 of 14
No
Yes
Drer\si:ch211-57n23.4
2 of 14
No
Yes
Drer\boc
1 of 14
No
No
Drer\cdon
1 of 14
No
No
Drer\chl1b
1 of 14
No
No
Drer\cntn3a.1
1 of 14
No
No
Drer\cntn3b
1 of 14
No
No
Drer\cntn4
1 of 14
No
No
Drer\cntn5
1 of 14
No
No
Drer\dscama
1 of 14
No
No
Drer\dscamb
1 of 14
No
No
Drer\dscaml1
1 of 14
No
No
Drer\fn1b
1 of 14
No
Yes
Drer\fndc1
1 of 14
No
Yes
Drer\hmcn1
1 of 14
No
No
Drer\hmcn2
1 of 14
No
No
Drer\hspg2
1 of 14
No
No
Drer\igsf9ba
1 of 14
No
No
Drer\igsf9b
1 of 14
No
No
Drer\lrrn1
1 of 14
No
No
Drer\myom1a
1 of 14
No
Yes
Drer\myot
1 of 14
No
Yes
Drer\mypn
1 of 14
No
Yes
Drer\ncam3
1 of 14
No
No
Drer\obscnb
1 of 14
No
No
Drer\obsl1b
1 of 14
No
No
Drer\palld
1 of 14
No
No
Drer\ptprdb
1 of 14
No
No
Drer\ptprd
1 of 14
No
No
Drer\ptprfa
1 of 14
No
No
Drer\ptprfb
1 of 14
No
No
Drer\ptprq
1 of 14
No
No
Drer\ptprsa
1 of 14
No
No
Drer\pxdn
1 of 14
No
No
Drer\robo1
1 of 14
No
No
Drer\robo2
1 of 14
No
No
Drer\robo3
1 of 14
No
No
Drer\sdk1a
1 of 14
No
No
Drer\sdk2a
1 of 14
No
No
Drer\sdk2b
1 of 14
No
No
Drer\ttn.1
1 of 14
No
No
Drer\ttn.2
1 of 14
No
No
Drer\unc5a
1 of 14
No
No
Drer\unc5da
1 of 14
No
No
Drer\zgc:77784
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (22)
Cele\unc-40
12 of 14
Yes
Yes
Cele\him-4
2 of 14
No
No
Cele\ptp-3
2 of 14
No
No
Cele\unc-89
2 of 14
No
No
Cele\clr-1
1 of 14
No
No
Cele\dig-1
1 of 14
No
Yes
Cele\ptp-4
1 of 14
No
No
Cele\pxn-1
1 of 14
No
No
Cele\rig-4
1 of 14
No
No
Cele\rig-5
1 of 14
No
No
Cele\sax-3
1 of 14
No
No
Cele\T13H5.1
1 of 14
No
No
Cele\unc-22
1 of 14
No
No
Cele\ZC374.2
1 of 14
No
Yes
Cele\zig-1
1 of 14
No
Yes
Cele\zig-2
1 of 14
No
No
Cele\zig-3
1 of 14
No
No
Cele\zig-4
1 of 14
No
No
Cele\zig-5
1 of 14
No
Yes
Cele\zig-7
1 of 14
No
Yes
Cele\zig-8
1 of 14
No
No
Cele\zig-11
1 of 14
No
Yes
Anopheles gambiae (African malaria mosquito) (70)
Agam\AgaP_AGAP006083
11 of 12
Yes
Yes
Agam\AgaP_AGAP000659
1 of 12
No
No
Agam\AgaP_AGAP000720
1 of 12
No
No
Agam\AgaP_AGAP000863
1 of 12
No
No
Agam\AgaP_AGAP001126
1 of 12
No
No
Agam\AgaP_AGAP001201
1 of 12
No
No
Agam\AgaP_AGAP001239
1 of 12
No
No
Agam\AgaP_AGAP001242
1 of 12
No
No
Agam\AgaP_AGAP001304
1 of 12
No
No
Agam\AgaP_AGAP002040
1 of 12
No
No
Agam\AgaP_AGAP002135
1 of 12
No
No
Agam\AgaP_AGAP002325
1 of 12
No
No
Agam\AgaP_AGAP002336
1 of 12
No
No
Agam\AgaP_AGAP002494
1 of 12
No
No
Agam\AgaP_AGAP002628
1 of 12
No
No
Agam\AgaP_AGAP002707
1 of 12
No
No
Agam\AgaP_AGAP002708
1 of 12
No
No
Agam\AgaP_AGAP002737
1 of 12
No
No
Agam\AgaP_AGAP002832
1 of 12
No
No
Agam\AgaP_AGAP002949
1 of 12
No
No
Agam\AgaP_AGAP003187
1 of 12
No
No
Agam\AgaP_AGAP003235
1 of 12
No
No
Agam\AgaP_AGAP003610
1 of 12
No
No
Agam\AgaP_AGAP004246
1 of 12
No
No
Agam\AgaP_AGAP004902
1 of 12
No
No
Agam\AgaP_AGAP004915
1 of 12
No
No
Agam\AgaP_AGAP005028
1 of 12
No
No
Agam\AgaP_AGAP005068
1 of 12
No
No
Agam\AgaP_AGAP005069
1 of 12
No
No
Agam\AgaP_AGAP005219
1 of 12
No
No
Agam\AgaP_AGAP005471
1 of 12
No
No
Agam\AgaP_AGAP005794
1 of 12
No
No
Agam\AgaP_AGAP006274
1 of 12
No
No
Agam\AgaP_AGAP006350
1 of 12
No
No
Agam\AgaP_AGAP007092
1 of 12
No
No
Agam\AgaP_AGAP007500
1 of 12
No
No
Agam\AgaP_AGAP007563
1 of 12
No
No
Agam\AgaP_AGAP007759
1 of 12
No
No
Agam\AgaP_AGAP008077
1 of 12
No
No
Agam\AgaP_AGAP008734
1 of 12
No
No
Agam\AgaP_AGAP008778
1 of 12
No
No
Agam\AgaP_AGAP008813
1 of 12
No
No
Agam\AgaP_AGAP009138
1 of 12
No
No
Agam\AgaP_AGAP009183
1 of 12
No
No
Agam\AgaP_AGAP009242
1 of 12
No
No
Agam\AgaP_AGAP009245
1 of 12
No
No
Agam\AgaP_AGAP009247
1 of 12
No
No
Agam\AgaP_AGAP009248
1 of 12
No
No
Agam\AgaP_AGAP009250
1 of 12
No
No
Agam\AgaP_AGAP009253
1 of 12
No
No
Agam\AgaP_AGAP009965
1 of 12
No
No
Agam\AgaP_AGAP010090
1 of 12
No
No
Agam\AgaP_AGAP010093
1 of 12
No
No
Agam\AgaP_AGAP010221
1 of 12
No
No
Agam\AgaP_AGAP010222
1 of 12
No
No
Agam\AgaP_AGAP010242
1 of 12
No
No
Agam\AgaP_AGAP010422
1 of 12
No
No
Agam\AgaP_AGAP010541
1 of 12
No
Yes
Agam\AgaP_AGAP010542
1 of 12
No
No
Agam\AgaP_AGAP010742
1 of 12
No
No
Agam\AgaP_AGAP010884
1 of 12
No
No
Agam\AgaP_AGAP010969
1 of 12
No
No
Agam\AgaP_AGAP010970
1 of 12
No
No
Agam\AgaP_AGAP011128
1 of 12
No
No
Agam\AgaP_AGAP011194
1 of 12
No
No
Agam\AgaP_AGAP011195
1 of 12
No
No
Agam\AgaP_AGAP012441
1 of 12
No
No
Agam\DOME
1 of 12
No
No
Agam\HPX4
1 of 12
No
No
Agam\IGL1
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:fra. 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) (62)
boi
2 of 13
ihog
2 of 13
Lar
2 of 13
plum
2 of 13
Pxn
2 of 13
sdk
2 of 13
Ama
1 of 13
btl
1 of 13
CG7166
1 of 13
CG34353
1 of 13
DIP-α
1 of 13
DIP-β
1 of 13
DIP-δ
1 of 13
DIP-ε
1 of 13
DIP-η
1 of 13
DIP-γ
1 of 13
DIP-ι
1 of 13
DIP-κ
1 of 13
DIP-λ
1 of 13
DIP-θ
1 of 13
DIP-ζ
1 of 13
dome
1 of 13
dpr1
1 of 13
dpr2
1 of 13
dpr3
1 of 13
dpr4
1 of 13
dpr5
1 of 13
dpr6
1 of 13
dpr7
1 of 13
dpr8
1 of 13
dpr9
1 of 13
dpr10
1 of 13
dpr11
1 of 13
dpr12
1 of 13
dpr13
1 of 13
dpr14
1 of 13
dpr15
1 of 13
dpr16
1 of 13
dpr17
1 of 13
dpr18
1 of 13
dpr19
1 of 13
dpr20
1 of 13
dpr21
1 of 13
Dscam1
1 of 13
Dscam2
1 of 13
Dscam3
1 of 13
Dscam4
1 of 13
et
1 of 13
klg
1 of 13
Lac
1 of 13
Nrg
1 of 13
otk
1 of 13
Ptp36E
1 of 13
Ptp69D
1 of 13
Ptp99A
1 of 13
robo1
1 of 13
robo2
1 of 13
robo3
1 of 13
sls
1 of 13
Strn-Mlck
1 of 13
trol
1 of 13
zormin
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Allele
Disease
Evidence
References
fra4
model of  Down syndrome
CEA with Dscam105518
(Hernández et al., 2023)
Potential Models Based on Orthology ( 3 )
Human Ortholog
Disease
Evidence
References
DCC; DCC netrin 1 receptor
model of  mirror movements 1
IEA
(FlyBase, 2019-)
model of  esophageal cancer
IEA
(FlyBase, 2019-)
model of  colorectal cancer
IEA
(FlyBase, 2019-)
Modifiers Based on Experimental Evidence ( 2 )
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?
DCC; DCC netrin 1 receptor
12 of 14
DCC NETRIN 1 RECEPTOR; DCC
 
NEO1; neogenin 1
12 of 14
601907
      IGDCC4; immunoglobulin superfamily DCC subclass member 4
      3 of 14
      616810
          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
          fra
          Hsap\DCC
          (Chaudhari et al., 2024)
          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
          fra
          enhanceable
          Abl::Hsap\ABL1::Hsap\BCR
          (Dorsten et al., 2010)
          fra
          enhanceable
          Abl
          (O'Donnell and Bashaw, 2013, Dorsten et al., 2010, Dorsten et al., 2007, Forsthoefel et al., 2005)
          fra
          suppressible
          Abl
          (Dorsten et al., 2007)
          fra
          enhanceable
          comm
          (Yang et al., 2009)
          fra
          suppressible
          comm
          (Yang et al., 2009)
          fra
          enhanceable
          Cyfip
          (Chaudhari et al., 2024)
          fra
          enhanceable
          drl
          (O'Donnell and Bashaw, 2013)
          fra
          enhanceable
          Dscam1
          (Raza and Jacobs, 2016)
          fra
          enhanceable
          Dscam1|Dscam3
          (Andrews et al., 2008)
          fra
          suppressible
          ena
          (Forsthoefel et al., 2005)
          fra
          suppressible
          fra::unc-5
          (Keleman and Dickson, 2001)
          fra
          enhanceable
          Iqsec
          (Hummel et al., 1999)
          fra
          enhanceable
          mew
          (Kolesnikov and Beckendorf, 2005)
          fra
          enhanceable
          mud
          (Cate et al., 2016)
          fra
          enhanceable
          mys
          (O'Donnell and Bashaw, 2013)
          fra
          enhanceable
          RhoGEF64C
          (Bashaw et al., 2001)
          fra
          enhanceable
          robo1
          (Raza and Jacobs, 2016, O'Donnell and Bashaw, 2013)
          fra
          suppressible
          robo1
          (de Torres-Jurado et al., 2022, Raza and Jacobs, 2016, Yang et al., 2009, Garbe et al., 2007)
          fra
          enhanceable
          robo2
          (Evans et al., 2015)
          fra
          enhanceable
          SCAR
          (Chaudhari et al., 2024)
          fra|Rho1
          suppressible
          sqh
          (Dorsten et al., 2007)
          fra
          suppressible
          Src42A
          (O'Donnell and Bashaw, 2013)
          fra
          suppressible
          Src42A|Src64B
          (O'Donnell and Bashaw, 2013)
          fra
          enhanceable
          Src64B
          (O'Donnell and Bashaw, 2013)
          fra
          suppressible
          Src64B
          (O'Donnell and Bashaw, 2013)
          fra
          enhanceable
          Tace
          (Zang et al., 2022)
          fra
          suppressible
          Tace
          (Zang et al., 2022)
          fra
          enhanceable
          trio
          (Forsthoefel et al., 2005)
          fra
          enhanceable
          tutl
          (Al-Anzi and Wyman, 2009)
          fra
          enhanceable
          unc-5
          (O'Donnell and Bashaw, 2013)
          Starting gene(s)
          Interaction type
          Interacting gene(s)
          Reference
          Abl::Hsap\ABL1::Hsap\BCR
          enhanceable
          fra
          (Dorsten et al., 2007)
          Abl::Hsap\ABL1::Hsap\BCR
          suppressible
          fra
          (Dorsten et al., 2010)
          Cdc42
          suppressible
          fra
          (Dorsten et al., 2007)
          Dscam1
          enhanceable
          fra
          (Raza and Jacobs, 2016)
          Dys
          suppressible
          fra
          (Kucherenko et al., 2008)
          Gαq
          suppressible
          fra
          (Hiramoto and Hiromi, 2006, Ratnaparkhi et al., 2002)
          Gαq
          enhanceable
          fra
          (Ratnaparkhi et al., 2002)
          Iqsec
          enhanceable
          fra
          (Onel et al., 2004, Hummel et al., 1999)
          N
          enhanceable
          fra
          (Kuzina et al., 2011)
          NetA
          suppressible
          fra
          (Winberg et al., 1998)
          NetB
          suppressible
          fra
          (Winberg et al., 1998)
          Rac1
          suppressible
          fra
          (Dorsten et al., 2007)
          Rac1
          enhanceable
          fra
          (Dorsten et al., 2007)
          Rho1
          enhanceable
          fra
          (Dorsten et al., 2007)
          robo1
          enhanceable
          fra
          (Raza and Jacobs, 2016)
          robo1
          suppressible
          fra
          (de Torres-Jurado et al., 2022, Raza and Jacobs, 2016, Gilestro, 2008, Garbe et al., 2007, Hiramoto and Hiromi, 2006)
          robo2
          suppressible
          fra
          (Raza and Jacobs, 2016)
          sli
          suppressible
          fra
          (Kolesnikov and Beckendorf, 2005)
          Tace
          enhanceable
          fra
          (Zang et al., 2022)
          Trim9
          suppressible
          fra
          (Morikawa et al., 2011, Song et al., 2011)
          Trim9
          enhanceable
          fra
          (Song et al., 2011)
          Trim9
          enhanceable
          robo1|fra
          (Song et al., 2011)
          unc-5
          suppressible
          fra
          (Keleman and Dickson, 2001)
          External Data
          Linkouts
          BioGRID - A database of protein and genetic interactions.
          DroID - A comprehensive database of gene and protein interactions.
          MIST (genetic) - An integrated Molecular Interaction Database
          MIST (protein-protein) - An integrated Molecular Interaction Database
          Pathways
          Signaling Pathways (FlyBase)
          Metabolic Pathways
          FlyBase
          External Links
          External Data
          Linkouts
          Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
          Class of Gene
          Genomic Location and Detailed Mapping Data
          Chromosome (arm)
          2R
          Recombination map
          2-67
          Cytogenetic map
          49A10-49B3
          Sequence location
          FlyBase Computed Cytological Location
          Cytogenetic map
          Evidence for location
          49A10-49B3
          Limits computationally determined from genome sequence between P{EP}CG8569EP1182 and P{EP}AmphEP2175&P{lacW}spt4k05316
          Experimentally Determined Cytological Location
          Cytogenetic map
          Notes
          References
          49B-49B
          (Timpe, 1996.9.16)
          49B-49B
          (determined by in situ hybridisation)
          (Kolodziej et al., 1996)
          Experimentally Determined Recombination Data
          Location

          2-67

          (FlyBase, 2016)

          2-69.0

          (Comeron et al., 2012)

          2-

          (Timpe, 1996.9.16)
          Left of (cM)
          Right of (cM)
          Notes
          Stocks and Reagents
          Stocks (71)
          Genomic Clones (31)
          cDNA Clones (75)
           

          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

          (Kolodziej et al., 1996)
          Commercially Available Antibodies
           
          Cell Line Information
          Publicly Available Cell Lines
           
            Other Stable Cell Lines
             
            Other Comments

            Expression of fra in R cells is not sufficient to alter target layer specificity.

            (Timofeev et al., 2012)

            fra is directly activated by en.

            (Joly et al., 2007)

            The growth cone response (attractive) is determined by the cytoplasmic domain of fra.

            (Keleman and Dickson, 2001)

            fra contributes to long range, but not short range, Netrin-mediated repulsion.

            (Keleman and Dickson, 2001)

            fra protein can capture Netrin protein (encoded by NetA and NetB) and "present" it for recognition by other receptors.

            (Hiramoto et al., 2000)

            The cytoplasmic domain of the guidance receptor fra can determine the growth cones' response in vivo, mediating attraction independently of the ectodomain and its particular ligand binding.

            (Bashaw and Goodman, 1999)

            The fra gene product is required in the target for attracting retinal fibers in the developing visual system. fra is not required for the differentiation of cells in the target.

            (Gong et al., 1999)

            One EMS induced allele was identified in a screen for mutations affecting commissure formation in the CNS of the embryo.

            (Hummel et al., 1999)

            fra is required for Netrin-mediated attraction, but not repulsion.

            (Winberg et al., 1998)

            fra encodes a DDC-related (deleted in colorectal cancer) protein. fra is expressed on axons in the developing nervous system as well as on a variety of epithelia. Null mutations in fra disrupt axon guidance in the central nervous system and motor axon pathfinding and target recognition in the periphery. fra functions cell autonomously raising the possibility that it encodes a receptor or part of a receptor mediating Netrin-dependent guidance.

            (Kolodziej et al., 1996)
            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 (12)
            Reported As
            Symbol Synonym
            CG8581
            (National Institute of Genetics Fly Stocks, 2022-, Ni et al., 2010.12.1, Schiess et al., 2009, Ni et al., 2008)
            CT24981
            (Hynes and Zhao, 2000)
            DCC
            (Yang et al., 2009, Hiramoto and Hiromi, 2006)
            Dcc
            (Richardson and Portela, 2018)
            Fra
            (Sanhueza et al., 2025, Owings and Chow, 2024, Gorla and Bashaw, 2020, Kang et al., 2019, Simonetti et al., 2019, Suzuki et al., 2018, Hughes and Jacobs, 2017, Gupta et al., 2016, Ozkan et al., 2013, Lu et al., 2012, Nose, 2012, Brierley et al., 2009, Gilestro, 2008, Talbot and VanBerkum, 2006, Murphey and Godenschwege, 2002)
            fra
            (Lopez et al., 2025, Sigalova et al., 2025, Sullivan and Bashaw, 2025, Chaudhari et al., 2024, Fischer et al., 2024, Lapraz et al., 2024, Sood et al., 2024, Okada et al., 2023, Xue et al., 2023, Clarembaux-Badell et al., 2022, Joseph et al., 2022, Zang et al., 2022, McLaughlin et al., 2021, Russell et al., 2021, Uçkun et al., 2021, Yin et al., 2021, Lecheta et al., 2020, Arbeille and Bashaw, 2018, Ros et al., 2018, Aw et al., 2017, Bonneaud et al., 2017, Cheong and VanBerkum, 2017, Hernandez-Fleming et al., 2017, Santiago and Bashaw, 2017, Cate et al., 2016, Evans, 2016, Halachmi et al., 2016, Raza and Jacobs, 2016, Shen et al., 2016, Doggett et al., 2015, Duff et al., 2015, Evans et al., 2015, Gene Disruption Project members, 2015-, Kok et al., 2015, Neuhaus-Follini and Bashaw, 2015, Gupta et al., 2014, Wolfram et al., 2014, Zarin et al., 2014, Czech et al., 2013, Das et al., 2013, Macabenta et al., 2013, Manavalan et al., 2013, Mora et al., 2013, Newquist et al., 2013, O'Donnell and Bashaw, 2013, O'Donnell and Bashaw, 2013, Ozkan et al., 2013, Ozkan et al., 2013, Busser et al., 2012, Timofeev et al., 2012, Kuzina et al., 2011, Matthews and Grueber, 2011, Morikawa et al., 2011, Song et al., 2011, Carrillo et al., 2010, Coleman et al., 2010, Dorsten et al., 2010, Flannery et al., 2010, Mrkusich et al., 2010, Spitzweck et al., 2010, von Hilchen et al., 2010, Al-Anzi and Wyman, 2009, Mauss et al., 2009, Tan et al., 2009, Yamamoto et al., 2009.2.25, Yang et al., 2009, Andrews et al., 2008, Dorsten and VanBerkum, 2008, Hsouna and VanBerkum, 2008, Kind et al., 2008, Kucherenko et al., 2008, O'Donnell et al., 2008, Ou et al., 2008, Bhat et al., 2007, Chen et al., 2007, Dorsten et al., 2007, Garbe and Bashaw, 2007, Garbe et al., 2007, Joly et al., 2007, Lecuyer et al., 2007, Simionato et al., 2007, Hiramoto and Hiromi, 2006, Pickeral et al., 2000)
            fra/DCC
            (Santiago and Bashaw, 2017)
            frazzled
            (Valdes-Aleman et al., 2021, Furrer et al., 2003)
            Name Synonyms
            DCC/Frazzled
            (Murphey and Godenschwege, 2002)
            Frazzled
            (Long et al., 2025, Warecki et al., 2025, Losada-Perez, 2018, Millard and Pecot, 2018, Richardson and Portela, 2018, Suzuki et al., 2018, Gupta et al., 2016, Neuhaus-Follini and Bashaw, 2015, Gupta et al., 2014, Zarin et al., 2014, Macabenta et al., 2013, O'Donnell and Bashaw, 2013, O'Donnell and Bashaw, 2013, Nose, 2012, Morikawa et al., 2011, Coleman et al., 2010, Dorsten et al., 2010, Al-Anzi and Wyman, 2009, Brierley et al., 2009, Tan et al., 2009, Dorsten and VanBerkum, 2008, Gilestro, 2008, Hsouna and VanBerkum, 2008, Kind et al., 2008, Tripodi et al., 2008, Dorsten et al., 2007, Mauss et al., 2007, Chilton, 2006, Talbot and VanBerkum, 2006)
            Netrin-receptor
            (Banyai and Patthy, 2004)
            deleted in colorectal cancer
            (Yang et al., 2009)
            frazzled
            (Kim et al., 2023, Chaturvedi and Murray, 2022, Mitchell, 2018, Hernandez-Fleming et al., 2017, Oliva et al., 2016, Manhire-Heath et al., 2013, Mora et al., 2013, Newquist et al., 2013, O'Donnell and Bashaw, 2013, Timofeev et al., 2012, Kuzina et al., 2011, Matthews and Grueber, 2011, Meyer et al., 2011, Mrkusich et al., 2010, von Hilchen et al., 2010, Liu et al., 2009, Mauss et al., 2009, Yang et al., 2009, Zlatic et al., 2009, Andrews and Kidd, 2008, Garbe et al., 2007, Joly et al., 2007, Hiramoto and Hiromi, 2006, Muraro and Moffat, 2006, Burnette et al., 2005, Furrer et al., 2003, Timpe, 1996.9.16, Timpe, 1996.9.16)
            Secondary FlyBase IDs
              Datasets (0)
              Study focus (0)
              Experimental Role
              Project
              Project Type
              Title
              Study result (0)
              Result
              Result Type
              Title
              External Crossreferences and Linkouts ( 56 )
              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.
              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
              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
              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.
              References (277)