FB2025_05 , released December 11, 2025
Gene: Dmel\brk
Open Close
General Information
Symbol
Dmel\brk
Species
D. melanogaster
Name
brinker
Annotation Symbol
CG9653
Feature Type
protein_coding_gene
FlyBase ID
FBgn0024250
Gene Model Status
Current
Stock Availability
26 publicly available
Gene Summary
brinker (brk) encodes a transcriptional repressor that plays important roles in the development of many tissues, including most imaginal discs, where its expression is negatively regulated by Dpp signaling. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Also Known As

ssg-1

Key Links
Genomic Location
Cytogenetic map
7B1-7B1
Sequence location
Recombination map
1-20
RefSeq locus
NC_004354 REGION:7307939..7311132
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 (10 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
enables DNA-binding transcription repressor activity, RNA polymerase II-specific
inferred from direct assay
(Weiss et al., 2010, Saller and Bienz, 2001)
enables RNA polymerase II transcription regulatory region sequence-specific DNA binding
inferred from direct assay
(Weiss et al., 2010, Stöbe et al., 2009, Kwon et al., 2004, Saller and Bienz, 2001, Sivasankaran et al., 2000)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (7 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
involved_in cell competition in a multicellular organism
inferred from mutant phenotype
(Moreno et al., 2002)
involved_in imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Dworkin and Gibson, 2006)
involved_in negative regulation of BMP signaling pathway
inferred from mutant phenotype
(Saller and Bienz, 2001, Torres-Vazquez et al., 2001, Campbell and Tomlinson, 1999, Minami et al., 1999)
inferred from direct assay
(Weiss et al., 2010)
involved_in negative regulation of transcription by RNA polymerase II
inferred from mutant phenotype
(Weiss et al., 2010, Kwon et al., 2004, Campbell and Tomlinson, 1999)
inferred from direct assay
(Saller and Bienz, 2001, Sivasankaran et al., 2000)
involved_in posterior Malpighian tubule development
inferred from mutant phenotype
(Hatton-Ellis et al., 2007)
involved_in regulation of gene expression
inferred from mutant phenotype
(Stöbe et al., 2009)
involved_in wing disc morphogenesis
inferred from mutant phenotype
(Moser and Campbell, 2005)
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
is_active_in nucleus
inferred from direct assay
(Ell et al., 2024)
located_in nucleus
inferred from direct assay
(Campbell and Tomlinson, 1999)
Terms Based on Predictions or Assertions (0 terms)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
brinker (brk) encodes a transcriptional repressor that plays important roles in the development of many tissues, including most imaginal discs, where its expression is negatively regulated by Dpp signaling. [Date last reviewed: 2019-03-07]
Gene Group (FlyBase)
UNCLASSIFIED DNA BINDING DOMAIN TRANSCRIPTION FACTORS -
This group comprises DNA-binding transcription factors that do not classify under other domain-based transcription factor groups in FlyBase.
Pathway (FlyBase)
NEGATIVE REGULATORS OF BMP SIGNALING PATHWAY -
Negative regulators of Bone Morphogenetic Protein (BMP) signaling down-regulate the pathway, ultimately resulting in the decreased nuclear activity of the Mad/Med transcription factor complex.
Summary (Interactive Fly)

novel transcription factor bearing weak homology to homeobox domain proteins - negatively regulates decapentaplegic target genes - pposing inputs by Hedgehog and Brinker define a stripe of hairy expression in the Drosophila leg imaginal discs - Mad, TCF and Brinker first activates then represses dpp expression in the posterior spiracles

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

Please see the JBrowse view of Dmel\brk 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 Q9XTN4)

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
PDB - An information portal to biological macromolecular structures
Comments on Gene Model

Gene model reviewed during 5.50

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
brk-RA
FBtr0071048
NM_078514
3194
704
Additional Transcript Data and Comments
Reported size (kB)

3.654 (longest cDNA)

(Campbell and Tomlinson, 1999)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
brk-PA
FBpp0071007
77.5
704
6.88
Q9XTN4
NP_511069
AAF46251
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)

704 (aa)

(Campbell and Tomlinson, 1999, Jazwinska et al., 1999)
Comments
External Data
Crossreferences
InterPro - A database of protein families, domains and functional sites
PDB - An information portal to biological macromolecular structures
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\brk 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).

-1.12

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
ectoderm | ventro-lateral
(Jazwinska et al., 1999)
embryonic stage 4 -- 5
ectoderm | ventro-lateral
(Zhang et al., 2001)
ectoderm | lateral | ventral
(Dunipace et al., 2013)
ectoderm
(Dunipace et al., 2013)
embryonic stage 4 -- 6
mesectoderm anlage
(Fisher et al., 2012)
ventral ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
embryonic stage 5
ventral neurectoderm
(Markstein et al., 2004)
(Dunipace et al., 2019)
embryonic stage 5 -- 6
mesectoderm
(Kearney et al., 2004)
embryonic stage 7 -- 8
central brain anlage
(Fisher et al., 2012)
head mesoderm
(Fisher et al., 2012)
mesectoderm
(Fisher et al., 2012)
trunk mesoderm
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
ventral nerve cord primordium

Comment: reported as ventral nerve cord anlage

(Fisher et al., 2012)
mesectoderm anlage
(Kearney et al., 2004)
embryonic stage 8 -- 9
ectoderm
(Dunipace et al., 2019)
embryonic stage 9
embryonic hindgut | dorsal
(Hatton-Ellis et al., 2007)
embryonic posterior Malpighian tubule
(Hatton-Ellis et al., 2007)
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)
mesectoderm
(Fisher et al., 2012)
salivary gland body primordium

Comment: reported as salivary gland body specific anlage

(Fisher et al., 2012)
trunk mesoderm
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
ventral nerve cord primordium
(Fisher et al., 2012)
embryonic stage 9 -- 11
ventral nerve cord primordium
(Jazwinska et al., 1999)
embryonic stage 9 -- 12
midline primordium
(Kearney et al., 2004)
embryonic stage 10
mesoderm | ventral
(Jazwinska et al., 1999)
embryonic hindgut | ventral
(Hamaguchi et al., 2012)
embryonic stage 11
tracheal pit | vicinity of
(Jazwinska et al., 1999)
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)
fat body/gonad primordium
(Fisher et al., 2012)
foregut primordium
(Fisher et al., 2012)
hindgut proper primordium
(Fisher et al., 2012)
lateral cord neuron
(Fisher et al., 2012)
midline primordium
(Fisher et al., 2012)
posterior midgut primordium
(Fisher et al., 2012)
salivary gland body primordium
(Fisher et al., 2012)
tracheal primordium
(Fisher et al., 2012)
ventral epidermis primordium
(Fisher et al., 2012)
embryonic stage 12
embryonic/larval midgut
(Jazwinska et al., 1999)
embryonic stage 13
parasegment 5 of endoderm
(Jazwinska et al., 1999)
visceral trunk mesoderm | restricted
(Jazwinska et al., 1999)
endoderm | restricted
(Jazwinska et al., 1999)
parasegment 5 of visceral trunk mesoderm
(Jazwinska et al., 1999)
embryonic stage 13 -- 16
embryonic dorsal epidermis
(Fisher et al., 2012)
embryonic head epidermis
(Fisher et al., 2012)
embryonic ventral epidermis
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
larval muscle system
(Fisher et al., 2012)
ventral midline of embryo
(Fisher et al., 2012)
larval midline neuron
(Kearney et al., 2004)
embryonic stage 14
ectoderm | ventral
(Jazwinska et al., 1999)
third instar larval stage
leg disc | restricted
(Jazwinska et al., 1999)
wing disc | posterior
(Campbell and Tomlinson, 1999)
wing disc | anterior
(Campbell and Tomlinson, 1999)
wing disc | restricted
(Jazwinska et al., 1999, Minami et al., 1999)
Additional Descriptive Data

brk is expressed in a narrow lateral stripe in early stage 5 embryos and in a broad lateral stripe at late stage 5. In stage 8-9 embryos, it is expressed in the ectoderm. Different enhancers are required for parts of the pattern.

(Dunipace et al., 2019)

At 4-h AEL, brk mRNA is expressed on the

(Hatton-Ellis et al., 2007)

dorsal side of the embryonic hindgut, and later is confined to the everting posterior tubule buds.

(Hatton-Ellis et al., 2007)

brk transcripts are expressed at the anterior and posterior extremes of the wing disc.

(Campbell and Tomlinson, 1999)

brk transcripts are expressed in lateral regions of the wing disc in a complementary pattern to that of dpp transcripts. In leg discs, brk expression is lowest in the dorsal compartment where dpp transcripts are expressed.

(Jazwinska et al., 1999)

In late syncytial and cellular blastoderm embryos, brk transcripts are expressed in a 9-10 nuclei-wide ventro-lateral stripe. The stripe expands to 18 nuclei wide shortly before gastrulation. A narrow gap initially separates the stripe dorsally from the dpp expression domain. Ventrally, the stripe is separated by one row of cells from the sna domain. The lateral brk domain largely overlaps the sog expression domain. During germ band extension, brk continues to be expressed in the ventral ectoderm where it appears to be restricted to neuroectodermal cells during neuroblast segregation. At stage 10, new sites of expression are observed in the ventral mesoderm. During stage 11, some ectodermal cells surrounding the tracheal pits express brk. brk expression is observed in the developing midgut during germ band retraction. In stage 13, three domains of brk expression in the visceral mesoderm and endoderm of the developing gut are observed. One is slightly anterior to the region of the developing gastric caecae, one is approximately at the location of parasegment 5, and one is at the border of the midgut and hindgut. During dorsal closure, brk transcripts are expressed in a broad stripe of ectodermal cells localized ventrally to the leading edge cells in which dpp controls morphogenesis. In general, brk is expressed in a complementary manner to regions of dpp signalling.

(Jazwinska et al., 1999)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 5
organism | restricted | ventro-lateral
(Helman et al., 2012)
third instar larval stage
wing disc | posterior
(Campbell and Tomlinson, 1999)
wing disc | anterior
(Campbell and Tomlinson, 1999)
antennal disc | dorsal | subset
(Firth et al., 2010)
eye disc | dorsal | subset
(Firth et al., 2010)
peripodial epithelium of eye disc
(Firth et al., 2010)
peripodial epithelium of antennal disc
(Firth et al., 2010)
Additional Descriptive Data

In antennal discs, brk protein is detected in the nuclei of cells in the dorsal antennal disc, except at the anterior-posterior compartment border, approximately reciprocal to the transcription of dpp just anterior to the compartment border, especially in the ventral antenna. In eye discs, brk protein is only detected in cells adjoining the dorsal anterior antenna, at the very anterior of the disc. Robust brk expression is detected in most cells of the peripodial epithelium overlying both eye and antennal discs.

(Firth et al., 2010)

brk protein is expressed at the anterior and posterior extremes of the wing disc. It is also detected in a gradient along the anterior-posterior axis in the wing pouch.

(Campbell and Tomlinson, 1999)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
is_active_in nucleus
inferred from direct assay
(Ell et al., 2024)
located_in nucleus
inferred from direct assay
(Campbell and Tomlinson, 1999)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\brk 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
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
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 ( 43 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 86 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of brk
Transgenic constructs containing regulatory region of brk
Aberrations (Deficiencies and Duplications) ( 11 )
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
cell lethal, with Scer\GAL4Ubx.PdC
lethal
lethal, with Scer\GAL4en-e16E
lethal (with brkKG08470)
lethal (with brkMcp)
lethal | recessive
lethal - all die before end of pupal stage | recessive
lethal - all die during embryonic stage | recessive
partially lethal
partially lethal, with Scer\GAL4en-e16E
partially lethal - majority die
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4en-e16E
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
Sterility
Allele
female fertile
female semi-fertile
Other Phenotypes
Allele
abnormal neuroanatomy
abnormal neuroanatomy, with Scer\GAL4Kr.PM
abnormal size | adult stage, with Scer\GAL4C-765
abnormal size | adult stage, with Scer\GAL4GMR.PF
abnormal size | adult stage, with Scer\GAL4nub-AC-62
abnormal size | adult stage, with Scer\GAL4vg.PU
abnormal size | larval stage, with Scer\GAL4nub-AC-62
decreased cell number, with Scer\GAL4hh-Gal4
decreased cell number, with Scer\GAL4nub-AC-62
decreased cell number | somatic clone, with Scer\GAL4FRT.Act5C
decreased occurrence of cell division | cell autonomous | somatic clone | third instar larval stage, with Scer\GAL4Act5C.Switch.PR
decreased occurrence of cell division | third instar larval stage, with Scer\GAL4hh-Gal4
decreased occurrence of cell division | third instar larval stage, with Scer\GAL4nub-AC-62
increased cell death, with Scer\GAL4hh-Gal4
increased cell death, with Scer\GAL4nub-AC-62
increased cell death | embryonic stage, with Scer\GAL4how-24B
increased cell death | somatic clone
increased cell death | somatic clone | third instar larval stage
increased cell number | adult stage, with Scer\GAL4esg.PU
increased cell number | somatic clone
increased occurrence of cell division | adult stage, with Scer\GAL4esg.PU
increased occurrence of cell division | cell non-autonomous | somatic clone | third instar larval stage, with Scer\GAL4Act5C.Switch.PR
increased occurrence of cell division | somatic clone | third instar larval stage
neoplasia | adult stage
neoplasia | somatic clone | third instar larval stage
some die during embryonic stage
some die during embryonic stage | recessive
some die during larval stage
some die during pupal stage | recessive
visible
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4C-765
visible, with Scer\GAL4en-e16E
visible, with Scer\GAL4nub-AC-62
visible | adult stage, with Scer\GAL4GMR.PF
visible | adult stage, with Scer\GAL4vg.PU
visible | adult stage | somatic clone
visible | cell autonomous | recessive | somatic clone
visible | cold sensitive, with Scer\GAL4Bx-MS1096
visible | dominant
visible | recessive
visible | somatic clone
Phenotype manifest in
Allele
abdominal 1 ventral denticle belt
adult midgut epithelium, with Scer\GAL4esg.PU
alar lobe
chrorion micropyle | somatic clone
crossvein | ectopic, with Scer\GAL4C-765
dorsal appendage | maternal effect | somatic clone
dorsal appendage | somatic clone
dorsal ectoderm
dorsal vessel wall cell, with Scer\GAL4how-24B, Scer\GAL4twi.PG
egg operculum | somatic clone
embryonic/first instar larval cuticle, with Scer\GAL4VP16.mat.αTub67C
embryonic/larval crystal cell
embryonic/larval crystal cell, with Scer\GAL4twi.PG
embryonic/larval epidermis
embryonic abdominal dorsal sensory organ precursor cluster, with Scer\GAL4Kr.PM
embryonic abdominal lateral sensory organ precursor cluster, with Scer\GAL4Kr.PM
embryonic abdominal ventral sensory organ precursor cluster group
embryonic epidermis
embryonic epidermis | dorsal
embryonic epidermis | ventral
embryonic head
embryonic primordium of adult salivary gland
embryonic primordium of adult salivary gland, with Scer\GAL4da.G32
embryonic ventral epidermis
eye, with Scer\GAL4GMR.PF
eye disc morphogenetic furrow | somatic clone, with Scer\GAL4Act5C.PI
hypostomal sclerite
imaginal disc, with Scer\GAL4nub-AC-62
larval dorsal hair
larval longitudinal connective
neuroblast
neuroblast NB2-4
neuroblast NB3-3
neuroblast NB4-2
neuroblast NB6-4
neuroblast NB7-3
neuroblast NB7-4
posterior crossvein, with Scer\GAL4en-e16E
posterior crossvein | heat sensitive, with Scer\GAL4en-e16E
presumptive embryonic/larval peripheral nervous system, with Scer\GAL4Kr.PM
salivary gland
symmetrical commissure
ventral denticle belt
ventral neurectoderm
wing
wing, with Scer\GAL4bi-omb-Gal4
wing, with Scer\GAL4Bx-MS1096
wing, with Scer\GAL4C-765
wing, with Scer\GAL4en-e16E
wing, with Scer\GAL4nub-AC-62
wing, with Scer\GAL4salm-459.2
wing, with Scer\GAL4vg.PU
wing | cell autonomous | somatic clone
wing | cold sensitive, with Scer\GAL4Bx-MS1096
wing | heat sensitive, with Scer\GAL4C-765
wing | lateral | somatic clone
wing | posterior, with Scer\GAL4en-e16E
wing | posterior | heat sensitive, with Scer\GAL4en-e16E
wing | somatic clone
wing blade, with Scer\GAL4C-765
wing disc, with Scer\GAL4en-e16E
wing disc, with Scer\GAL4Ubx.PdC
wing disc | cell autonomous | third instar larval stage
wing disc | cell non-autonomous | medial | third instar larval stage
wing disc | lateral | somatic clone
wing disc | somatic clone
wing disc | somatic clone, with Scer\GAL4FRT.Act5C
wing disc posterior compartment, with Scer\GAL4hh-Gal4
wing margin | heat sensitive, with Scer\GAL4en-e16E
wing vein, with Scer\GAL4Bx-MS1096
wing vein, with Scer\GAL4C-765
wing vein, with Scer\GAL4en-e16E
wing vein, with Scer\GAL4nub-AC-62
wing vein, with Scer\GAL4salm-459.2
wing vein, with Scer\GAL4vg.int2.1
wing vein | ectopic
wing vein | ectopic, with Scer\GAL4C-765
wing vein | ectopic | somatic clone
wing vein | somatic clone
wing vein L4, with Scer\GAL4en-e16E
wing vein L4 | heat sensitive, with Scer\GAL4en-e16E
wing vein L5, with Scer\GAL4en-e16E
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) (1)
Hsap\DAXX
1 of 14
Yes
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) (1)
Rnor\Daxx
1 of 14
Yes
No
Mus musculus (laboratory mouse) (1)
Mmus\Daxx
1 of 14
Yes
No
Xenopus tropicalis (Western clawed frog) (1)
Xtro\daxx
1 of 13
Yes
No
Danio rerio (Zebrafish) (1)
Drer\daxx
1 of 14
Yes
No
Caenorhabditis elegans (Nematode, roundworm) (0)
Anopheles gambiae (African malaria mosquito) (2)
Agam\AgaP_AGAP010938
7 of 12
Yes
Yes
Agam\AgaP_AGAP009432
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:brk. 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) (2)
CG14441
1 of 13
Daxx
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?
    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
    brk
    suppressible
    Abd-B
    (Kyrchanova and Georgiev, 2010)
    brk
    suppressible
    Dad
    (Martin et al., 2004)
    brk|yki|tkv
    enhanceable
    Mad
    (Oh and Irvine, 2011)
    brk
    suppressible
    mir-ban
    (Oh and Irvine, 2011)
    brk
    suppressible
    Myc
    (Doumpas et al., 2013)
    brk
    suppressible
    pan
    (Yang et al., 2013)
    brk
    suppressible
    puc
    (Moreno et al., 2002)
    brk
    suppressible
    shn
    (Torres-Vazquez et al., 2001)
    brk
    suppressible
    tkv
    (Torres-Vazquez et al., 2001)
    brk
    suppressible
    tkv|Mad
    (Oh and Irvine, 2011)
    brk
    enhanceable
    tsh
    (Saller et al., 2002)
    brk
    suppressible
    yki
    (Oh and Irvine, 2011)
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    ey
    suppressible
    brk
    (Kango-Singh et al., 2003)
    nab
    suppressible
    brk
    (Suissa et al., 2011)
    shn
    suppressible
    brk
    (Rusten et al., 2002, Torres-Vazquez et al., 2001, Marty et al., 2000)
    tkv
    suppressible
    brk
    (Martin et al., 2004, Torres-Vazquez et al., 2001)
    tsh
    enhanceable
    brk
    (Saller et al., 2002)
    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
    SignaLink - A signaling pathway resource with multi-layered regulatory networks.
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    X
    Recombination map
    1-20
    Cytogenetic map
    7B1-7B1
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    7B1-7B1
    Limits computationally determined from genome sequence between P{EP}CG9650EP1340&P{EP}CG9650EP1617 and P{EP}CBPEP1643&P{EP}EP1523EP1523
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    7A-7A
    (Campbell, 1999.2.4)
    7B-7B
    (determined by in situ hybridisation)
    (Minami et al., 1999)
    7B1-7B1
    (Jazwinska et al., 1999)
    Experimentally Determined Recombination Data
    Location

    1-20

    (FlyBase, 2016)

    1-21.0

    (Comeron et al., 2012)

    1-

    (Jazwinska et al., 1998, Rushlow et al., 1998)

    1-19.6

    (Rushlow, 1998.3.17)
    Left of (cM)
    (Jazwinska et al., 1999)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (26)
    Genomic Clones (10)
     

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

    cDNA Clones (22)
     

    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

    (Doumpas et al., 2013, Dunipace et al., 2013, Yang et al., 2013, Campbell and Tomlinson, 1999)
    Commercially Available Antibodies
     
    Cell Line Information
    Publicly Available Cell Lines
     
      Other Stable Cell Lines
       

      • New stable cell line derived from S2-unspecified : 75 stable S2 cell lines containing a library of intrinsically disordered regions (IDRs) fluorescently tagged with mNeonGreen were constructed. In addition, stable S2 cell lines expressing the following full-length transcription factors were generated: da, rib, CG10321, CG13287, tgo, Spps, CG31510, brk, and disco.

        (Hannon and Eisen, 2024)
      Other Comments

      The brk product negatively regulates dac expression in the embryonic head.

      (Anderson et al., 2006)

      The brk silencer serves as a direct target for a protein complex consisting of Mad/Med and shn.

      (Muller et al., 2003)

      3 alleles of brk been recovered in a screen for mutations with mutant phenotypes in clones in the wing.

      (Vegh and Basler, 2003)

      The brk co-repressors gro and CtBP are alternately used for repressing some dpp-responsive genes, whereas for repressing other, distinct, target genes they are not required.

      (Hasson et al., 2001)

      brk not only plays a role in dorsal-ventral but also in anterior-posterior axis patterning.

      (Lammel et al., 2000)

      dpp can control gene expression indirectly by downregulating the expression of brk, which represses dpp targets. Overall orientation not stated: brk- anon-7Ab+

      (Campbell and Tomlinson, 1999)

      The medial-to-lateral dpp gradient along the anterior-posterior axis in the developing wing is complemented by a lateral-to-medial brk gradient, and the opposition of these two gradients may allow cells to detect small differences in dpp concentration and respond by activating different target genes.

      (Campbell and Tomlinson, 1999)

      The embryonic phenotype of brk mutants indicates an involvement in dpp signalling.

      (Jazwinska et al., 1999)

      brk acts as a dpp antagonist in wing patterning. brk function is most important in regions where the dpp gradient has diminishing levels or where a further spreading of the dpp signal has to be prevented.

      (Jazwinska et al., 1999)

      brk represses dpp target genes independently of dl in the early embryo. brk has differential effects on dpp target genes, depending on the level of dpp activity required for their activation.

      (Jazwinska et al., 1999)

      brk protein negatively regulates dpp-dependent genes.

      (Minami et al., 1999)

      brk mutations are embryonic lethal. Homozygous embryos show an expansion of dorsal epidermis at the expense of ventral epidermis. Mosaic analysis indicates that brk is required for imaginal disc patterning.

      (Jazwinska et al., 1998)

      brk acts cell-autonomously as a negative element within the dpp pathway, acting downstream of dpp.

      (Jazwinska et al., 1998)

      brk mutant embryos show an embryonic dorsal-ventral phenotype where the denticle belts are reduced in width. brk also affects adult appendage development and wing vein patterning. Increasing the number of copies of brk+ in male flies enhances the dpp wing vein phenotype. The dpp wing vein phenotype is suppressed in female flies carrying a single copy of brk+.

      (Rushlow et al., 1998)
      Relationship to Other Genes
      Source for database merge of
      Additional comments

      "l(1)G0082" may affect "brk". "l(1)G0182" may affect "brk". "l(1)G0029" may affect "brk".

      (Peter et al., 2002)
      Nomenclature History
      Source for database identify of
      Nomenclature comments
      Etymology
      Synonyms and Secondary IDs (9)
      Reported As
      Symbol Synonym
      BRK
      (Blatti et al., 2015, Itasaki and Hoppler, 2010)
      Brk
      (Yusupova and Fuchs, 2023, Montanari et al., 2022, Rossi et al., 2021, Sharifkhodaei and Auld, 2021, Chen and Desplan, 2020, Gou et al., 2020, Wu et al., 2019, Ruiz-Losada et al., 2018, Bosch et al., 2017, Romero-Pozuelo et al., 2017, Zaytseva and Quinn, 2017, Matsuda et al., 2016, Mbodj et al., 2016, Bier and De Robertis, 2015, Harmansa et al., 2015, Esteves et al., 2014, Doumpas et al., 2013, Mbodj et al., 2013, Morozov and Ioshikhes, 2013, Turki-Judeh and Courey, 2012, Hamaratoglu et al., 2011, Oh and Irvine, 2011, Schwank et al., 2011, Quijano et al., 2010, Guruharsha et al., 2009, Stöbe et al., 2009, Campbell and Moser, 2008, Lembong et al., 2008, Liang et al., 2007, Tyler et al., 2007, Ashe, 2006, Cordier et al., 2006, Angulo et al., 2004, Song et al., 2004, Markstein et al., 2002, Xu et al., 2001)
      CG9653
      (Ni et al., 2011.7.19, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Perkins et al., 2009, Grieder et al., 2007, Legube et al., 2006, Blanchette et al., 2004, Bourbon et al., 2002)
      Dm brk
      (Crocker et al., 2008)
      brinker
      (Costa-Rodrigues et al., 2021)
      brk
      (El Marzkioui et al., 2025, Rasouliha et al., 2025, Tesson and Vincent, 2025, Al Asafen et al., 2024, Ell et al., 2024, Hunt and Mannervik, 2024, Malin et al., 2024, Adelmann et al., 2023, Floc'hlay et al., 2023, Matsuda and Affolter, 2023, Roth, 2023, Akiyama et al., 2022, Beaver et al., 2022, Chen et al., 2022, Dunipace et al., 2022, Romanova-Michaelides et al., 2022, Valencia-Expósito et al., 2022, Vuilleumier et al., 2022, Kvon et al., 2021, Matsuda et al., 2021, McParland et al., 2021, Ruiz-Losada et al., 2021, Strassburger et al., 2021, Zhao et al., 2021, Bakker et al., 2020, Berndt et al., 2020, Bosch et al., 2020, Haudry et al., 2020, Port et al., 2020, Schloop et al., 2020, Stathopoulos and Newcomb, 2020, Bredesen and Rehmsmeier, 2019, Chayengia et al., 2019, Chen, 2019, Gui et al., 2019, Romero-Pozuelo et al., 2019, Shokri et al., 2019, Bischof et al., 2018, Eusebio et al., 2018, Sander et al., 2018, Setiawan et al., 2018, Barrio and Milán, 2017, Hu et al., 2017.6.13, Karaiskos et al., 2017, Martín et al., 2017, Osterfield et al., 2017, Transgenic RNAi Project members, 2017-, Upadhyay et al., 2017, Dominguez et al., 2016, Esposito et al., 2016, Kwon et al., 2016, Norman et al., 2016, Sarov et al., 2016, Charbonnier et al., 2015, Denes et al., 2015, Luo et al., 2015, Merino et al., 2015, Schertel et al., 2015, Foo et al., 2014, Hodar et al., 2014, Mannervik, 2014, Navarro et al., 2014, Restrepo et al., 2014, Roy et al., 2014, Chen et al., 2013, Curtis et al., 2013, Dunipace et al., 2013, Gafner et al., 2013, Harbison et al., 2013, Hevia and de Celis, 2013, Humphreys et al., 2013, Li and Gilmour, 2013, Li et al., 2013, McKay and Lieb, 2013, Organista and De Celis, 2013, Saunders et al., 2013, Upadhyai and Campbell, 2013, Webber et al., 2013, Akiyama et al., 2012, Chopra et al., 2012, Hamaguchi et al., 2012, Helman et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Le and Wharton, 2012, Liang et al., 2012, Rebeiz et al., 2012, Rushlow and Shvartsman, 2012, Spokony and White, 2012.5.22, Stultz et al., 2012, Ziv et al., 2012, Abruzzi et al., 2011, Dworkin et al., 2011, Garcia and Stathopoulos, 2011, Hamaratoglu et al., 2011, Lott et al., 2011, Mrinal et al., 2011, Nien et al., 2011, Oh and Irvine, 2011, Ozdemir et al., 2011, Rodriguez, 2011, Schwank et al., 2011, Suissa et al., 2011, Thomas et al., 2011, Yang and Su, 2011, Firth et al., 2010, Popodi et al., 2010-, Portela et al., 2010, Schwank and Basler, 2010, Venken et al., 2010, Vuilleumier et al., 2010, Weiss et al., 2010, Yagi et al., 2010, Bai et al., 2009, Cai and Laughon, 2009, Foronda et al., 2009, Lembong et al., 2009, McKay et al., 2009, McNamee and Brodsky, 2009, Papatsenko et al., 2009, Song et al., 2009, Twombly et al., 2009, Widmann and Dahmann, 2009, Wolfstetter et al., 2009, Ziv et al., 2009, Zúñiga et al., 2009, Bates et al., 2008, Beaver et al., 2008, Bickel et al., 2008, Brás-Pereira and Casares, 2008, Charbonnier and Pyrowolakis, 2008, Estella and Mann, 2008, Estella et al., 2008, Fromental-Ramain et al., 2008, Fu and Levine, 2008, Hong et al., 2008, McNamee and Brodsky, 2008, Mieszczanek et al., 2008, Newfeld et al., 2008, Qi et al., 2008, Rogulja et al., 2008, Schwank et al., 2008, Warrior et al., 2008, Yao et al., 2008, Aerts et al., 2007, Barrio et al., 2007, Engström et al., 2007, Estella and Mann, 2007, Gao and Laughon, 2007, Gerlitz et al., 2007, Grieder et al., 2007, Hatton-Ellis et al., 2007, Li et al., 2007, Shravage et al., 2007, Theisen et al., 2007, Umemori et al., 2007, Walsh and Carroll, 2007, Zeitlinger et al., 2007, Zeitlinger et al., 2007, Anderson et al., 2006, Bangi and Wharton, 2006, Bangi and Wharton, 2006, Chen and Schupbach, 2006, Dworkin and Gibson, 2006, Gao and Laughon, 2006, Glinka et al., 2006, Herranz et al., 2006, Mizutani et al., 2006, Mosimann et al., 2006, Ratnaparkhi et al., 2006, Sotillos and de Celis, 2006, Stultz et al., 2006, Yao et al., 2006, Gao et al., 2005, Mehlen et al., 2005, Rogulja and Irvine, 2005, Sotillos and De Celis, 2005, Stathopoulos and Levine, 2005, Markstein et al., 2004, Pyrowolakis et al., 2004, Saller and Bienz, 2001)
      ssg-1
      (Lammel and Saumweber, 2000, Lammel et al., 2000)
      Name Synonyms
      Brinker
      (Collins and Anderson, 2018, Zaytseva and Quinn, 2017, Kwon et al., 2016, Saadin and Starz-Gaiano, 2016, Hariharan, 2015, Mannervik, 2014, Levayer and Moreno, 2013, Peterson and O'Connor, 2013, Liang et al., 2012, Eivers et al., 2011, Hamaratoglu et al., 2011, Quijano et al., 2010, Rhiner et al., 2010, Weiss et al., 2010, Campbell and Moser, 2008, Dougherty et al., 2008, Fromental-Ramain et al., 2008, Lembong et al., 2008, Newfeld et al., 2008, Rogulja et al., 2008, Takaesu et al., 2008, Winkler et al., 2008, Gerlitz et al., 2007, Hatton-Ellis et al., 2007, Liang et al., 2007, Cordier et al., 2006, Sotillos and De Celis, 2005, Song et al., 2004, Song et al., 2003, Markstein et al., 2002, Saller and Bienz, 2001, Xu et al., 2001)
      brinker
      (Cohen et al., 2020, Papadopoulos et al., 2019, Osterfield et al., 2017, Esposito et al., 2016, Charbonnier et al., 2015, Dunipace et al., 2013, Gafner et al., 2013, Harbison et al., 2013, McKay and Lieb, 2013, Peterson and O'Connor, 2013, Akiyama et al., 2012, Rebeiz et al., 2012, Lott et al., 2011, Oh and Irvine, 2011, Qin et al., 2011, Firth et al., 2010, Portela et al., 2010, Vuilleumier et al., 2010, Yagi et al., 2010, Bai et al., 2009, Cai and Laughon, 2009, Cande et al., 2009, Foronda et al., 2009, Krejcí et al., 2009, McNamee and Brodsky, 2009, Wolfstetter et al., 2009, Bickel et al., 2008, Brás-Pereira and Casares, 2008, Cook et al., 2008, Crocker et al., 2008, Fu and Levine, 2008, Hong et al., 2008, Qi et al., 2008, Schwank et al., 2008, Warrior et al., 2008, Yao et al., 2008, Estella and Mann, 2007, Fu and Levine, 2007, Gao and Laughon, 2007, Li et al., 2007, Umemori et al., 2007, Walsh and Carroll, 2007, Anderson et al., 2006, Dworkin and Gibson, 2006, Dworkin and Gibson., 2006, Mosimann et al., 2006, Sotillos and de Celis, 2006, Tsuda et al., 2006, Mehlen et al., 2005, Markstein et al., 2004, Campbell, 1999.2.4, Jazwinska et al., 1998, Rushlow et al., 1998)
      small salivary glands-1
      (Lammel et al., 2000)
      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 ( 40 )
        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)
        PDB - An information portal to biological macromolecular structures
        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.
        FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
        Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
        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
        SignaLink - A signaling pathway resource with multi-layered regulatory networks.
        References (468)