FB2020_06, released Dec 22, 2020

Gene: Dmel\brat

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General Information
Symbol
Dmel\brat
Species
D. melanogaster
Name
brain tumor
Annotation Symbol
CG10719
Feature Type
protein_coding_gene
FlyBase ID
FBgn0010300
Gene Model Status
Current
Stock Availability
39 publicly available
Gene Snapshot
brain tumor (brat) encodes a tumor suppressor that regulates proliferation in the brain. During neural stem cell division, it segregates into one daughter cell where it inhibits protein translation, preventing self-renewal and inducing differentiation. It also acts during oogenesis where it interacts with the products of nos and pum to suppress hb translation. [Date last reviewed: 2019-03-07]
Other Summaries
Also Known As

l(2)37Cf, lincRNA.S1565, l(2)E60

Key Links
Genomic Location
Cytogenetic map
37C1-37C6
Sequence location
2L:19,133,816..19,179,758 [+]
Recombination map

2-54

RefSeq locus
NT_033779 REGION:19133816..19179758
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
GO Summary Ribbons
Gene Ontology (GO) Annotations (33 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
enables mRNA 3'-UTR binding
inferred from mutant phenotype
(Reichardt et al., 2018)
enables protein binding
inferred from physical interaction with UniProtKB:P25724
(assigned by UniProt )
(Sonoda and Wharton, 2001)
inferred from physical interaction with UniProtKB:Q8MQJ9
(assigned by UniProt )
(Sonoda and Wharton, 2001)
inferred from physical interaction with FLYBASE:eIF4EHP; FB:FBgn0053100
(Cho et al., 2006)
enables translation repressor activity
inferred from mutant phenotype
(assigned by UniProt )
(Sonoda and Wharton, 2001)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
enables mRNA 3'-UTR binding
inferred from biological aspect of ancestor with PANTHER:PTN004670338
(assigned by GO_Central )
(Gaudet et al., 2011)
enables translation regulator activity
traceable author statement
(Dean et al., 2002)
enables translation repressor activity
inferred from biological aspect of ancestor with PANTHER:PTN000647790
(assigned by GO_Central )
(Gaudet et al., 2011)
enables zinc ion binding
inferred from electronic annotation with InterPro:IPR000315
(assigned by InterPro )
(FlyBase Curators et al., 2004-)
Biological Process (22 terms)
Terms Based on Experimental Evidence (21 terms)
CV Term
Evidence
References
involved_in asymmetric cell division
inferred from mutant phenotype
(Betschinger et al., 2006)
involved_in asymmetric neuroblast division resulting in ganglion mother cell formation
inferred from mutant phenotype
(Lee et al., 2006)
involved_in brain development
inferred from mutant phenotype
(Bello et al., 2006)
involved_in ganglion mother cell fate determination
inferred from mutant phenotype
(Liu et al., 2017, Betschinger et al., 2006)
involved_in negative regulation of cell population proliferation
inferred from mutant phenotype
(Bello et al., 2006)
involved_in negative regulation of gene expression
inferred from mutant phenotype
(Landskron et al., 2018, Reichardt et al., 2018)
involved_in negative regulation of neuroblast proliferation
inferred from mutant phenotype
(Betschinger et al., 2006)
inferred from mutant phenotype
(assigned by UniProt )
(Janssens et al., 2014)
inferred from genetic interaction with UniProtKB:Q9VQ56
(assigned by UniProt )
(Janssens et al., 2014)
involved_in negative regulation of translation
inferred from mutant phenotype
(Reichardt et al., 2018)
involved_in neuroblast development
inferred from mutant phenotype
(assigned by UniProt )
(Bowman et al., 2008)
involved_in neuroblast differentiation
inferred from mutant phenotype
(Reichardt et al., 2018)
involved_in neuroblast fate determination
inferred from mutant phenotype
(Komori et al., 2014)
involved_in neuroblast proliferation
inferred from mutant phenotype
(assigned by UniProt )
(Bowman et al., 2008)
inferred from mutant phenotype
(Caussinus and Gonzalez, 2005)
involved_in neuron differentiation
inferred from mutant phenotype
(Reichardt et al., 2018, Lee et al., 2006)
involved_in oogenesis
inferred from high throughput mutant phenotype
(Schupbach and Wieschaus, 1991)
involved_in regulation of neuroblast proliferation
inferred from genetic interaction with UniProtKB:P25439
(assigned by UniProt )
(Janssens et al., 2014)
inferred from genetic interaction with UniProtKB:Q26263
(assigned by UniProt )
(Janssens et al., 2014)
involved_in regulation of neurogenesis
inferred from mutant phenotype
(Zhu et al., 2011)
involved_in regulation of synaptic growth at neuromuscular junction
inferred from mutant phenotype
(Shi et al., 2013)
involved_in regulation of synaptic vesicle endocytosis
inferred from mutant phenotype
(Shi et al., 2013)
involved_in rRNA transcription
inferred from mutant phenotype
(assigned by UniProt )
(Frank et al., 2002)
involved_in segmentation
inferred from mutant phenotype
(Cho et al., 2006)
involved_in ventral cord development
inferred from high throughput mutant phenotype
(Bivik et al., 2015)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
involved_in negative regulation of translation
inferred from biological aspect of ancestor with PANTHER:PTN000647790
(assigned by GO_Central )
(Gaudet et al., 2011)
involved_in regulation of translation
traceable author statement
(Dean et al., 2002)
Cellular Component (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
located_in apical cortex
inferred from direct assay
(Betschinger et al., 2006)
located_in basal cortex
inferred from direct assay
(Betschinger et al., 2006)
located_in cell cortex region
inferred from mutant phenotype
(Reichardt et al., 2018)
located_in cytoplasm
inferred from direct assay
(assigned by UniProt )
(Sonoda and Wharton, 2001)
located_in cytosol
inferred from direct assay
(Betschinger et al., 2006)
located_in neuronal cell body
inferred from direct assay
(Shi et al., 2013)
Terms Based on Predictions or Assertions (0 terms)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Protein Function (UniProtKB)
A NHL-domain family protein that functions a translational repressor to inhibit cell proliferation (PubMed:11274060). Plays a central role in translation repression of hb mRNA by being recruited by nos and pum to the Nanos Response Element (NRE), a 16 bp sequence in the hb mRNA 3'-UTR (PubMed:11274060). Probably recruited by other proteins to repress translation of other mRNAs in other tissues (PubMed:11274060). Involved in the regulation of ribosomal RNA synthesis and cell growth (PubMed:11807032). Participates in abdominal segmentation and imaginal disk development (PubMed:11274060). During neuroblast division, segregates asymmetrically and inhibits self-renewal of one of the two daughter cells (PubMed:16564014). Together with the asymmetrically segregating transcription factor prospero ensures that the daughter cell will stop growing, exit the cell cycle, and differentiate into neurons possibly by modulating the function of dm in ganglion mother cells (GMC) (PubMed:11807032, PubMed:16564014). Restrics developmental potential of type II intermediary neuronal progenitor (INP) cells playing a role in proliferation and maturation of the neuroblasts (PubMed:24550111, PubMed:14561773, PubMed:22143802, PubMed:18342578).
(UniProt, Q8MQJ9)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
l(2)37Cf
Lethality occurs as pharate adults. Third-instar larvae of 13 of 14 alleles examined display enlarged brains-up to eight times normal volume; greatest in presumptive optic center (Hankins and Wright). Brain fragments transplanted into normal adults show uncontrolled proliferation and metastatic invasion of other tissues (Gateff). Escapers [l(2)37Cf1/l(2)37Cfts3] lethargic and sterile in both sexes [Wright, 1987, Results and Problems in Cell Differentiation (W. Hennig, ed.). Springer-Verlag, Berlin, Heidelberg, pp. 95-120]. Homozygous females [l(2)37Cffs2] retain eggs in ovaries (Schupbach).
Summary (Interactive Fly)
Gene Model and Products
Number of Transcripts
5
Number of Unique Polypeptides
2

Please see the JBrowse view of Dmel\brat for information on other features

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

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Comments on Gene Model

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

Gene model reviewed during 5.48

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.45

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

Multiple (sequential) stage-specific extensions of 3' UTRs observed during embryogenesis (FBrf0215804); all variants may not be annotated.

Gene model reviewed during 5.55

Sequence Ontology: Class of Gene
nuclear_gene
protein_coding_gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
brat-RA
FBtr0081158
NM_057597
4544
1037
brat-RB
FBtr0081159
NM_134302
4963
1037
brat-RC
FBtr0081160
NM_206004
12088
1037
brat-RE
FBtr0336627
NM_001201913
6258
1061
brat-RF
FBtr0344846
NM_001299154
4869
1037
Additional Transcript Data and Comments
Reported size (kB)

4.933 (longest cDNA)

(Arama et al., 2000)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
brat-PA
FBpp0080702
110.4
1037
7.21
NP_476945
AAN11027
brat-PB
FBpp0080703
110.4
1037
7.21
NP_599129
AAF53771
brat-PC
FBpp0089239
110.4
1037
7.21
NP_995726
AAS64719
brat-PE
FBpp0307610
113.2
1061
7.30
NP_001188842
ADV37091
brat-PF
FBpp0311161
110.4
1037
7.21
NP_001286083
AHN54597
Polypeptides with Identical Sequences

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

1037 aa isoforms: brat-PA, brat-PB, brat-PC, brat-PF
Additional Polypeptide Data and Comments
Reported size (kDa)

1037 (aa)

(Arama et al., 2000)
Comments
External Data
Subunit Structure (UniProtKB)

Interacts with nos and pum (PubMed:11274060). Acts via the formation of a quaternary complex composed of pum, nos, brat and the 3'-UTR mRNA of hb (PubMed:11274060). Not recruited by nos and pum to cyclin B 3'-UTR mRNA (PubMed:11274060). Might interact with mira; the interaction seems to be important for brat localization during mitosis (PubMed:16564014).

(UniProt, Q8MQJ9)
Domain

The NHL repeats form six-bladed beta-propeller that mediate the interaction with the pumilio repeats of pum, and are essential for translational effector function.

(UniProt, Q8MQJ9)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
UniProt
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\brat 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
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 16
peripheral nervous system precursor cluster
(Arama et al., 2000)
antenno-maxillary complex
(Arama et al., 2000)
embryonic stage 12 -- 17
embryonic ventral nervous system

Comment: expression starts late in embryonic stage 12

(Arama et al., 2000)
presumptive embryonic/larval peripheral nervous system

Comment: expression starts late in embryonicstage 12

(Arama et al., 2000)
embryonic brain

Comment: expression starts late in embryonic stage 12

(Arama et al., 2000)
embryonic stage 11 -- 17
(Arama et al., 2000)
embryonic stage 13 -- 16
embryonic ganglion mother cell
(Arama et al., 2000)
embryonic brain
(Fisher et al., 2012)
embryonic/larval nervous system
(Fisher et al., 2012)
ventral nerve cord
(Fisher et al., 2012)
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
central brain anlage in statu nascendi
(Fisher et al., 2012)
dorsal ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
mesoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
ventral ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
antennal anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

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

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)
visual anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(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)
organism | ubiquitous
(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)
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)
ventral nerve cord primordium
(Fisher et al., 2012)
embryonic stage 11 -- 12
central brain primordium
(Fisher et al., 2012)
embryonic central brain glial cell
(Fisher et al., 2012)
embryonic central brain neuron
(Fisher et al., 2012)
lateral cord glial cell
(Fisher et al., 2012)
lateral cord neuron
(Fisher et al., 2012)
ventral nerve cord primordium
(Fisher et al., 2012)
oogenesis
germarium
(Harris et al., 2011)
female germline stem cell
(Harris et al., 2011)
border follicle cell
(Wang et al., 2006)
embryonic stage 10 -- 15
embryonic/larval central nervous system

Comment: extended 3' UTR isoform

(Hilgers et al., 2011)
Additional Descriptive Data

brat transcripts are expressed throughout the germline, including in the germline stem cells, however brat protein is excluded from the germline stem cells.

(Harris et al., 2011)

brat transcripts are detected early and late in embryogenesis but transcripts containing the 3' UTR extension (~8.5 kb) are only detected at later stages and appear after the maternal-to-zygotic transition. Sequential, phased, 3' UTR lengthening is seen (3' UTR lengthening ocurring in multiple steps during later stages of embryogenesis). brat extended 3' UTR transcripts are expressed specifically in the nervous system.

(Hilgers et al., 2011)

brat expression is enriched in border follicle cells relative to other cells in the egg chamber.

(Wang et al., 2006)

brat is expressed in the developing brain, the ventral CNS, and the PNS starting at stage 11. Expression is detected in the ganglion mother cells starting at stage 13. In stage 16 embryos, 4 clusters of peripheral neurons express brat. The lacZ reporter construct Ecol\lacZbrat-k06028 shows additional staining in cardioblasts from stage 11 to 17, which was not detected by brat in situ hybridization. See Ecol\lacZbrat-k06028RA for reporter construct expression, including expression in larval stages.

(Arama et al., 2000)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
wandering third instar larval stage
motor neuron
(Shi et al., 2013)
ventral nerve cord | restricted
(Shi et al., 2013)
abdominal dorsal multidendritic neuron ddaC
(Olesnicky et al., 2012)
Additional Descriptive Data

While brat transcripts are expressed in germline stem cells, brat protein is excluded from germline stem cells.

(Harris et al., 2011)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in apical cortex
inferred from direct assay
(Betschinger et al., 2006)
located_in basal cortex
inferred from direct assay
(Betschinger et al., 2006)
located_in cell cortex region
inferred from mutant phenotype
(Reichardt et al., 2018)
located_in cytoplasm
inferred from direct assay
(assigned by UniProt )
(Sonoda and Wharton, 2001)
located_in cytosol
inferred from direct assay
(Betschinger et al., 2006)
located_in neuronal cell body
inferred from direct assay
(Shi et al., 2013)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\brat in GBrowse 2
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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
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, and Transgenic Constructs
Classical and Insertion Alleles ( 94 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 40 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of brat
Transgenic constructs containing regulatory region of brat
Deletions and Duplications ( 37 )
Disrupted in
Df(2L)BSC341
(Kahsai et al., 2016)
Df(2L)E71
(Brittnacher and Ganetzky, 1983, Wright et al., 1981)
Df(2L)hk39
(Brittnacher and Ganetzky, 1983)
Df(2L)NST
(Wright, 1996, Stathakis et al., 1995, Pentz et al., 1986)
Df(2L)Sd57
(Wright, 1996, Stathakis et al., 1995, Brittnacher and Ganetzky, 1983)
Df(2L)TE37C-1
(Wright, 1996, Gilbert et al., 1984, Wright et al., 1981)
Df(2L)TE37C-7
(Cho et al., 2006, Wright, 1996, Stathakis et al., 1995)
Df(2L)TW50
(Spradling et al., 1999, BDGP Project Members, 1994-1999, Schupbach and Wieschaus, 1991, Wright et al., 1976)
Df(2L)TW130
(Wright, 1996, Stathakis et al., 1995, Schupbach and Wieschaus, 1991, Gilbert et al., 1984, Brittnacher and Ganetzky, 1983, Wright et al., 1981, Wright et al., 1976)
Df(2L)TW158
(Brittnacher and Ganetzky, 1983)
Df(2L)VA12
(Wright, 1996, Stathakis et al., 1995, BDGP Project Members, 1994-1999, Pentz et al., 1986, Gilbert et al., 1984, Wright et al., 1981)
Df(2L)VA13
(Wright, 1996, Gilbert et al., 1984, Wright et al., 1981)
Df(2L)VA14
(Brittnacher and Ganetzky, 1983)
Df(2L)VA16
(Brittnacher and Ganetzky, 1983)
Df(2L)VA17
(Wright, 1996, Stathakis et al., 1995, BDGP Project Members, 1994-1999, Pentz et al., 1986, Gilbert et al., 1984, Wright et al., 1981)
Df(2L)VA19
(Wright, 1996, Stathakis et al., 1995, BDGP Project Members, 1994-1999, Brittnacher and Ganetzky, 1983)
Df(2L)VA20
(Brittnacher and Ganetzky, 1983)
Df(2L)VA23
(Stathakis et al., 1995)
Not disrupted in
Df(2L)E55
(Stathakis et al., 1995, Brittnacher and Ganetzky, 1983)
Df(2L)hk18
(Wright, 1996, Pentz et al., 1986, Gilbert et al., 1984)
Df(2L)hk-UC1
(Wright, 1996, Stathakis et al., 1995)
Df(2L)hk-UC2
(Wright, 1996, Stathakis et al., 1995)
Df(2L)M36F-S5
(BDGP Project Members, 1994-1999)
Df(2L)OD15
(Wright, 1996, Stathakis et al., 1995, BDGP Project Members, 1994-1999, Pentz et al., 1986)
Df(2L)pr26
(Brittnacher and Ganetzky, 1983)
Df(2L)Sd2
(Brittnacher and Ganetzky, 1983)
Df(2L)Sd14
(Brittnacher and Ganetzky, 1983)
Df(2L)Sd37
(Wright, 1996, Stathakis et al., 1995, Brittnacher and Ganetzky, 1983)
Df(2L)Sd77
(Stathakis et al., 1995, BDGP Project Members, 1994-1999, Brittnacher and Ganetzky, 1983)
Df(2L)TW2
(Brittnacher and Ganetzky, 1983)
Df(2L)TW9
(BDGP Project Members, 1994-1999)
Df(2L)TW137
(Stathakis et al., 1995, BDGP Project Members, 1994-1999)
Df(2L)TW203
(Wright, 1996, Stathakis et al., 1995, Pentz et al., 1986)
Df(2L)VA6
(Brittnacher and Ganetzky, 1983)
Df(2L)VA8
(Brittnacher and Ganetzky, 1983)
Df(2L)VA18
(Wright, 1996, Stathakis et al., 1995, BDGP Project Members, 1994-1999, Pentz et al., 1986, Gilbert et al., 1984)
Df(2L)VA21
(Wright, 1996, Stathakis et al., 1995)
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal
lethal, with Scer\GAL4insc-Mz1407
lethal (with Df(2L)BSC341)
lethal | recessive
lethal | recessive | heat sensitive
lethal - all die before end of embryonic stage | germline clone | maternal effect | recessive
lethal - all die before end of embryonic stage | germline clone | non-rescuable maternal effect | recessive
partially lethal - majority die, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
partially lethal - majority die | recessive
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4tin.CΔ4
Sterility
Allele
female sterile
female sterile, with Scer\GAL4VP16.nos.UTR
female sterile | recessive
fertile
Other Phenotypes
Allele
cell cycle defective, with Scer\GAL4da.PU
cell number defective | larval stage
cell number defective | third instar larval stage
decreased cell number, with Scer\GAL4VP16.nos.UTR
decreased cell number | cell non-autonomous, with Scer\GAL4en-e16E
decreased cell number | embryonic stage | maternal effect
decreased cell number | somatic clone, with Scer\GAL4Act5C.PP
endocytosis defective | third instar larval stage (with brat2L-150-11)
endocytosis defective | third instar larval stage (with brat2L-192-9)
endocytosis defective | third instar larval stage (with brat11)
increased cell number (with brat14)
increased cell number (with brat18)
increased cell number | germline clone
increased cell number | larval stage, with Scer\GAL4insc-Mz1407
increased cell number | larval stage | somatic clone
increased cell number | somatic clone | third instar larval stage
increased cell number | third instar larval stage
increased cell number | third instar larval stage, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
increased cell size, with Scer\GAL4dpp.blk1
increased cell size | germline clone
increased cell size | somatic clone
increased cell size | somatic clone, with Scer\GAL4Act5C.PP
increased cell size | somatic clone | third instar larval stage
increased occurrence of cell division | adult stage | somatic clone
increased occurrence of cell division | larval stage | somatic clone
increased occurrence of cell division | somatic clone | third instar larval stage
increased occurrence of cell division | third instar larval stage
mitotic cell cycle defective
neuroanatomy defective
neuroanatomy defective | larval stage
neuroanatomy defective | larval stage (with brat11)
neuroanatomy defective | larval stage (with bratDG19310)
neuroanatomy defective | larval stage (with bratk06028)
neuroanatomy defective | larval stage | somatic clone
neuroanatomy defective | somatic clone
neuroanatomy defective | somatic clone | third instar larval stage
neuroanatomy defective | third instar larval stage
neuroanatomy defective | third instar larval stage, with Scer\GAL4elav.PU
neuroanatomy defective | third instar larval stage (with brat2L-150-11)
neuroanatomy defective | third instar larval stage (with brat2L-192-9)
neuroanatomy defective | third instar larval stage (with brat11)
neuroanatomy defective | third instar larval stage (with Df(2L)pr-A16)
neurophysiology defective | third instar larval stage (with brat2L-150-11)
neurophysiology defective | third instar larval stage (with brat2L-192-9)
neurophysiology defective | third instar larval stage (with brat11)
oxidative stress response defective, with Scer\GAL4da.PU
short lived, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
size defective, with Scer\GAL4da.PU
size defective | adult stage, with Scer\GAL4nub.PU
some die during embryonic stage | maternal effect | recessive
some die during embryonic stage | non-rescuable maternal effect | recessive
tumorigenic
tumorigenic, with Scer\GAL4Act5C.PI
tumorigenic, with Scer\GAL4da.PU
tumorigenic, with Scer\GAL4insc-Mz1407
tumorigenic, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
tumorigenic (with brat11)
tumorigenic (with brat14)
tumorigenic (with brat18)
tumorigenic | adult stage, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
tumorigenic | larval stage, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
tumorigenic | somatic clone
tumorigenic | third instar larval stage
visible, with Scer\GAL4ey.PH
visible | cell non-autonomous, with Scer\GAL4en-e16E
wild-type
wild-type, with Scer\GAL4en-e16E
Phenotype manifest in
Allele
1st posterior cell, with Scer\GAL4dpp.blk1
adult brain, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
adult brain | somatic clone
adult thorax (with brat1)
adult thorax (with brat7)
apterous-expressing neuron of the lateral cluster of the thorax | ectopic
apterous-expressing neuron of the lateral cluster of the thorax Ap4 | supernumerary
axon | somatic clone
brain
brain, with Scer\GAL4insc-Mz1407
brain, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
egg chamber, with Scer\GAL4VP16.nos.UTR
embryo | germline clone | maternal effect | posterior
embryonic/larval brain
embryonic/larval brain, with Scer\GAL4Act5C.PI
embryonic/larval brain, with Scer\GAL4da.PU
embryonic/larval brain, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
embryonic/larval brain (with brat14)
embryonic/larval brain (with brat18)
embryonic/larval brain | cell autonomous | somatic clone
embryonic/larval brain | posterior
embryonic/larval brain | third instar larval stage
embryonic/larval neuromuscular junction | third instar larval stage, with Scer\GAL4Toll-6-D42
embryonic abdomen
embryonic abdomen (with brat1)
embryonic abdomen (with bratfs1)
embryonic abdomen (with bratfs3)
embryonic abdomen (with bratk06028)
embryonic abdomen (with Df(2L)TE37C-7)
embryonic abdominal segment
embryonic abdominal segment (with brat1)
embryonic abdominal segment (with bratfs1)
embryonic abdominal segment (with bratfs3)
embryonic abdominal segment (with bratk06028)
embryonic abdominal segment (with Df(2L)TE37C-7)
embryonic abdominal segment | maternal effect (with Df(2L)TE37C-7)
embryonic epidermis
embryonic epidermis (with brat1)
embryonic epidermis (with bratfs1)
embryonic epidermis (with bratfs3)
embryonic epidermis (with bratk06028)
embryonic epidermis (with Df(2L)TE37C-7)
eye, with Scer\GAL4ey.PH
female germline cell, with Scer\GAL4VP16.nos.UTR
female germline cyst, with Scer\GAL4VP16.nos.UTR
female germline stem cell, with Scer\GAL4VP16.nos.UTR
female germline stem cell | germline clone | supernumerary
ganglion mother cell
germarium, with Scer\GAL4VP16.nos.UTR
imaginal disc (with brat14)
imaginal disc (with brat18)
larval neuroblast | larval stage
mitotic cell cycle | somatic clone, with Scer\GAL4Act5C.PP
neuroblast, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
neuroblast | pupal stage, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
neuroblast | somatic clone
neuroblast | supernumerary
neuron
NMJ bouton | supernumerary | third instar larval stage, with Scer\GAL4elav.PU
NMJ bouton | supernumerary | third instar larval stage (with brat2L-150-11)
NMJ bouton | supernumerary | third instar larval stage (with brat2L-192-9)
NMJ bouton | supernumerary | third instar larval stage (with brat11)
NMJ bouton | supernumerary | third instar larval stage (with Df(2L)pr-A16)
NMJ bouton | third instar larval stage, with Scer\GAL4Toll-6-D42
nucleolus | somatic clone
postsynaptic density (with brat2L-192-9)
postsynaptic density (with brat11)
presynaptic active zone | third instar larval stage, with Scer\GAL4Toll-6-D42
presynaptic active zone membrane (with brat2L-192-9)
presynaptic active zone membrane (with brat11)
RP2 motor neuron | maternal effect
RP2 motor neuron | supernumerary, with Scer\GAL4VP16.mat.αTub67C
synaptic vesicle (with brat2L-192-9)
synaptic vesicle (with brat11)
T neuron T4 | somatic clone | third instar larval stage
T neuron T5 | somatic clone | third instar larval stage
type II neuroblast | larval stage | somatic clone | supernumerary
type II neuroblast | larval stage | supernumerary
type II neuroblast | larval stage | supernumerary, with Scer\GAL4insc-Mz1407
type II neuroblast | larval stage | supernumerary (with brat11)
type II neuroblast | larval stage | supernumerary (with bratDG19310)
type II neuroblast | larval stage | supernumerary (with bratk06028)
type II neuroblast | supernumerary
type II neuroblast | supernumerary (with brat11)
type II neuroblast | supernumerary (with bratDG19310)
type II neuroblast | third instar larval stage, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
wing, with Scer\GAL4nub.PU
wing & trichogen cell
wing blade anterior compartment | cell non-autonomous, with Scer\GAL4en-e16E
Orthologs
Downloads
Download All DIOPT Orthologs
Download All OrthoDB Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (12)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Hsap\TRIM3
7 of 15
Yes
Yes
Hsap\TRIM2
6 of 15
No
Yes
Hsap\NHLRC3
2 of 15
No
Yes
Hsap\NHLRC1
1 of 15
No
Yes
Hsap\RNF152
1 of 15
No
Yes
Hsap\RNF207
1 of 15
No
Yes
Hsap\RNF208
1 of 15
No
Yes
Hsap\TRIM25
1 of 15
No
Yes
Hsap\TRIM32
1 of 15
No
Yes
Hsap\TRIM45
1 of 15
No
No
Hsap\TRIM56
1 of 15
No
No
Hsap\TRIM71
1 of 15
No
No
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (11)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Mmus\Trim3
7 of 15
Yes
Yes
Mmus\Trim2
6 of 15
No
Yes
Mmus\Nhlrc3
2 of 15
No
Yes
Mmus\Nhlrc1
1 of 15
No
Yes
Mmus\Rnf207
1 of 15
No
Yes
Mmus\Rnf208
1 of 15
No
Yes
Mmus\Trim25
1 of 15
No
Yes
Mmus\Trim32
1 of 15
No
Yes
Mmus\Trim45
1 of 15
No
No
Mmus\Trim56
1 of 15
No
No
Mmus\Trim71
1 of 15
No
No
Rattus norvegicus (Norway rat) (10)
Rnor\Trim3
5 of 13
Yes
Yes
Rnor\Nhlrc3
2 of 13
No
Yes
Rnor\Trim2
2 of 13
No
No
Rnor\Nhlrc1
1 of 13
No
No
Rnor\Rnf152
1 of 13
No
Yes
Rnor\Rnf207
1 of 13
No
Yes
Rnor\Rnf208
1 of 13
No
Yes
Rnor\Trim32
1 of 13
No
Yes
Rnor\Trim45
1 of 13
No
No
Rnor\Trim71
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (8)
Xtro\trim2
5 of 12
Yes
Yes
Xtro\trim3
4 of 12
No
Yes
Xtro\nhlrc3
2 of 12
No
Yes
Xtro\rnf152
1 of 12
No
Yes
Xtro\rnf207
1 of 12
No
Yes
Xtro\trim32
1 of 12
No
Yes
Xtro\trim45
1 of 12
No
No
Xtro\trim71
1 of 12
No
No
Danio rerio (Zebrafish) (14)
Drer\trim2a
6 of 15
Yes
Yes
Drer\trim3b
6 of 15
Yes
Yes
Drer\trim3a
5 of 15
No
Yes
Drer\trim2b
4 of 15
No
Yes
Drer\ftr51
1 of 15
No
Yes
Drer\ftr85
1 of 15
No
Yes
Drer\rnf152
1 of 15
No
Yes
Drer\rnf207a
1 of 15
No
Yes
Drer\rnf207b
1 of 15
No
Yes
Drer\si:dkey-106l3.6
1 of 15
No
Yes
Drer\trim25
1 of 15
No
Yes
Drer\trim32
1 of 15
No
Yes
Drer\trim45
1 of 15
No
No
Drer\trim71
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (9)
Cele\ncl-1
12 of 15
Yes
Yes
Cele\lin-41
2 of 15
No
No
Cele\nhl-1
2 of 15
No
No
Cele\nhl-3
2 of 15
No
Yes
Cele\F40G9.12
1 of 15
No
Yes
Cele\F40G9.19
1 of 15
No
Yes
Cele\F43C11.8
1 of 15
No
No
Cele\F43G6.8
1 of 15
No
Yes
Cele\nhl-2
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (3)
Atha\AT1G56500
1 of 9
Yes
No
Atha\AT4G01350
1 of 9
Yes
Yes
Atha\emb1974
1 of 9
Yes
Yes
Saccharomyces cerevisiae (Brewer's yeast) (1)
Scer\PSH1
1 of 15
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG091903EZ )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila suzukii
Spotted wing Drosophila
Dsuz\DS10_00001265
Drosophila simulans
Dsim\GD24188
Drosophila sechellia
Dsec\GM17330
Drosophila erecta
Dere\GG21163
Drosophila yakuba
Dyak\GE13236
Drosophila ananassae
Dana\GF14706
Drosophila pseudoobscura pseudoobscura
Dpse\GA10519
Drosophila persimilis
Dper\GL21191
Drosophila willistoni
Dwil\GK23851
Drosophila virilis
Dvir\GJ18246
Drosophila mojavensis
Dmoj\GI13993
Drosophila grimshawi
Dgri\GH13439
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091501N8 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Mdoa\17011154
Glossina morsitans
Tsetse fly
Gmor\GMOY002193
Lucilia cuprina
Australian sheep blowfly
Lcup\FF38_07298
Mayetiola destructor
Hessian fly
Mdes\Mdes010620
Aedes aegypti
Yellow fever mosquito
Aaeg\AAEL004918
Anopheles darlingi
American malaria mosquito
Adar\ADAC005204
Anopheles gambiae
Malaria mosquito
Agam\AGAP010054
Culex quinquefasciatus
Southern house mosquito
Cqui\CPIJ017329
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W01YF )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bmor\BGIBMGA009425
Danaus plexippus
Monarch butterfly
Dple\DPOGS208423-PA
Heliconius melpomene
Postman butterfly
Hmel\HMEL012881
Apis florea
Little honeybee
Aflo\4336935348
Apis mellifera
Western honey bee
Amel\brat
Bombus impatiens
Common eastern bumble bee
Bimp\BIMP16815
Bombus terrestris
Buff-tailed bumblebee
Bter\15206974
Linepithema humile
Argentine ant
Lhum\LH16343
Megachile rotundata
Alfalfa leafcutting bee
Mrot\MROTU:002669
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG004513
Dendroctonus ponderosae
Mountain pine beetle
Dpod\YQE_02382
Dendroctonus ponderosae
Mountain pine beetle
Dpod\YQE_01340
Tribolium castaneum
Red flour beetle
Tcas\brat
Pediculus humanus
Human body louse
Phum\PHUM061260
Rhodnius prolixus
Kissing bug
Rpro\RPRC007254
Cimex lectularius
Bed bug
Clec\CLEC008353
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI007272
Zootermopsis nevadensis
Nevada dampwood termite
Znev\Znev_11025
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X01W2 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Smar\SMAR002957
Ixodes scapularis
Black-legged tick
Isca\ISCW011089
Stegodyphus mimosarum
African social velvet spider
Smim\KFM59712.1
Stegodyphus mimosarum
African social velvet spider
Smim\KFM61352.1
Tetranychus urticae
Two-spotted spider mite
Turt\tetur23g01840
Daphnia pulex
Water flea
Dpux\DAPPUDRAFT_299638
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G02UY )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_2527
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_966
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_638
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_1848
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_1135
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_1106
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_946
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_1093
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_2232
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_1027
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_665
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_902
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_438
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_401
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_567
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-trim2-4
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_2167
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_187
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-trim56L
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_354
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_1776
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-trim33l
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Hypp_846
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (5)
Gene Symbol
Score
Source
Compara
eggNOG
Homologene
Inparanoid
Isobase
OrthoDB
OrthoFinder
orthoMCL
Panther
RoundUp
Alignment
tn
4 of 10
wech
4 of 10
CG8419
2 of 10
mei-P26
2 of 10
Trim9
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Allele
Disease
Evidence
References
bratHMS01121
model of  cancer
CEA
(Hussain et al., 2017)
Potential Models Based on Orthology ( 1 )
Human Ortholog
Disease
Evidence
References
TRIM2; tripartite motif containing 2
model of  Charcot-Marie-Tooth disease type 2R
IEA
(FlyBase, 2019-)
Modifiers Based on Experimental Evidence ( 3 )
Disease Associations of Human Orthologs (via DIOPT v8.0 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?
TRIM3; tripartite motif containing 3
7 of 15
605493
      TRIM2; tripartite motif containing 2
      6 of 15
      614141
      Functional Complementation Data
      Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
      Interactions
      Summary of Physical Interactions
      esyN Network Diagram
      Show neighbor-neighbor interactions:
      Select Layout:
      Legend:
      Protein
      RNA
      Selected Interactor(s)
      Interactions Browser
      View in Interactions Browser
      Please see the Physical Interaction reports below for full details
      protein-protein
      Physical Interaction
      Assay
      References
      brat
      x-ray crystallography
      (Liu et al., 2019)
      brat - AGO1
      anti bait coimmunoprecipitation, western blot, two hybrid, anti tag coimmunoprecipitation, anti tag western blot, bimolecular fluorescence complementation, fluorescence microscopy, peptide massfingerprinting
      (Malik et al., 2017, Neumuller et al., 2008)
      brat - Sxl
      two hybrid, bimolecular fluorescence complementation, fluorescence microscopy, anti tag coimmunoprecipitation, anti tag western blot
      (Malik et al., 2017)
      brat - Tis11
      anti tag coimmunoprecipitation, anti tag western blot
      (Komori et al., 2018)
      brat - bam
      bimolecular fluorescence complementation, fluorescence microscopy, two hybrid, anti tag coimmunoprecipitation, anti tag western blot
      (Malik et al., 2017)
      brat - eIF4EHP
      anti tag coimmunoprecipitation, anti tag western blot, anti bait coimmunoprecipitation, western blot
      (Cho et al., 2006)
      brat - mira
      molecular sieving, molecular weight estimation by staining, two hybrid, isothermal titration calorimetry, predetermined participant, pull down, anti tag coimmunoprecipitation, anti tag western blot
      (Liu et al., 2019, Jia et al., 2015, Komori et al., 2014, Lee et al., 2006)
      brat - nos
      two hybrid
      (Sonoda and Wharton, 2001)
      brat - pum
      pull down, western blot, anti tag western blot, three hybrid, molecular sieving, molecular weight estimation by staining, transcriptional complementation assay
      (Loedige et al., 2014, Kadyrova et al., 2007, Cho et al., 2006, Edwards et al., 2001, Sonoda and Wharton, 2001)
      brat - sstn
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Liu et al., 2015)
      brat - stau
      pull down, molecular weight estimation by staining, anti tag coimmunoprecipitation, primer specific pcr
      (Jia et al., 2015, Loedige et al., 2015)
      RNA-protein
      Physical Interaction
      Assay
      References
      brat - Corin
      anti tag coimmunoprecipitation, primer specific pcr
      (Loedige et al., 2015)
      brat - Mad
      anti tag coimmunoprecipitation, quantitative reverse transcription pcr
      (Malik et al., 2019)
      brat - Myc
      anti tag coimmunoprecipitation, primer specific pcr
      (Loedige et al., 2015)
      brat - Pc
      anti tag coimmunoprecipitation, primer specific pcr
      (Loedige et al., 2015)
      brat - Sirt1
      anti tag coimmunoprecipitation, primer specific pcr
      (Loedige et al., 2015)
      brat - aub
      iclip, partial RNA sequence identification
      (Ma et al., 2017)
      brat - chinmo
      anti tag coimmunoprecipitation, primer specific pcr
      (Loedige et al., 2015)
      brat - dpn
      anti tag coimmunoprecipitation, primer specific pcr, electrophoretic mobility shift assay, autoradiography
      (Reichardt et al., 2018, Loedige et al., 2015)
      brat - elav
      anti bait coimmunoprecipitation, primer specific pcr
      (Hilgers et al., 2012)
      brat - hb
      anti tag coimmunoprecipitation, quantitative reverse transcription pcr, molecular sieving, static light scattering, nucleic acid uv cross-linking assay, Identification by mass spectrometry, microscale thermophoresis, predetermined participant, electrophoretic mobility shift assay, autoradiography
      (Malik et al., 2019, Loedige et al., 2014)
      brat - klu
      electrophoretic mobility shift assay, autoradiography, anti tag coimmunoprecipitation, primer specific pcr
      (Loedige et al., 2015)
      brat - kni
      electrophoretic mobility shift assay, autoradiography, anti tag coimmunoprecipitation, primer specific pcr
      (Loedige et al., 2015)
      brat - orb2
      pull down, western blot
      (Mastushita-Sakai et al., 2010)
      brat - par-6
      anti tag coimmunoprecipitation, primer specific pcr
      (Loedige et al., 2015)
      brat - piwi
      anti bait coimmunoprecipitation, northern blot
      (Saito et al., 2009)
      brat - stau
      pull down, molecular weight estimation by staining, anti tag coimmunoprecipitation, primer specific pcr
      (Jia et al., 2015, Loedige et al., 2015)
      brat - w
      anti tag coimmunoprecipitation, primer specific pcr
      (Loedige et al., 2015)
      brat - zld
      electrophoretic mobility shift assay, autoradiography, anti bait coimmunoprecipitation, quantitative reverse transcription pcr
      (Reichardt et al., 2018)
      Summary of Genetic Interactions
      esyN Network Diagram
      esyN Network Key:
      Suppression
      Enhancement
      View in Interactions Browser
      Please look at the allele data for full details of the genetic interactions
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      brat
      enhanceable
      Apc2
      (Komori et al., 2014)
      Apc2|brat
      suppressible
      arm
      (Komori et al., 2014)
      brat
      enhanceable
      arm
      (Komori et al., 2014)
      brat
      enhanceable
      ato
      (Mora et al., 2018)
      brat
      enhanceable
      Axn
      (Komori et al., 2014)
      brat
      enhanceable
      brm
      (Janssens et al., 2014)
      brat
      suppressible
      bsk
      (Hussain et al., 2017)
      brat
      enhanceable
      bsk
      (Hussain et al., 2017)
      brat
      suppressible
      btd
      (Komori et al., 2014)
      brat
      enhanceable
      CkIα
      (Komori et al., 2014)
      brat
      suppressible
      dpn
      (Reichardt et al., 2018)
      brat
      enhanceable
      dpn
      (Janssens et al., 2014)
      brat
      enhanceable
      E(spl)mγ-HLH
      (Janssens et al., 2014)
      brat
      suppressible
      E(spl)mγ-HLH
      (Janssens et al., 2014)
      brat
      enhanceable
      erm
      (Janssens et al., 2014)
      brat
      suppressible
      erm
      (Janssens et al., 2014)
      brat
      suppressible
      Gpdh1
      (Hussain et al., 2017)
      brat
      suppressible
      grh
      (Reichardt et al., 2018)
      brat
      suppressible
      Imp
      (Landskron et al., 2018)
      brat
      suppressible
      klu
      (Xiao et al., 2012)
      brat
      enhanceable
      klu
      (Janssens et al., 2014)
      brat
      suppressible
      lncRNA:cherub
      (Landskron et al., 2018)
      brat
      suppressible
      Mad
      (Shi et al., 2013)
      brat
      suppressible
      Mdh1
      (Hussain et al., 2017)
      brat
      suppressible
      Men
      (Hussain et al., 2017)
      brat
      suppressible
      Mfe2
      (Hussain et al., 2017)
      brat
      enhanceable
      Mfe2
      (Hussain et al., 2017)
      brat
      enhanceable
      mor
      (Janssens et al., 2014)
      brat
      suppressible
      Myc
      (Reichardt et al., 2018)
      brat
      suppressible
      orb2
      (Santana and Casas-Tintó, 2017)
      brat
      enhanceable
      osa
      (Janssens et al., 2014)
      brat
      suppressible
      Pepck1
      (Hussain et al., 2017)
      brat
      suppressible
      pros
      (Bello et al., 2006)
      brat
      enhanceable
      pros
      (Betschinger et al., 2006)
      brat
      enhanceable
      sgg
      (Komori et al., 2014)
      brat
      suppressible
      Syp
      (Landskron et al., 2018)
      brat
      enhanceable
      Syp
      (Landskron et al., 2018)
      brat
      suppressible
      trx
      (Komori et al., 2014)
      brat
      suppressible
      Wwox
      (Hussain et al., 2017)
      brat
      suppressible
      zld
      (Reichardt et al., 2018)
      brat
      suppressible
      Zw
      (Hussain et al., 2017)
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      ct
      suppressible
      brat
      (Krupp et al., 2005)
      gcm
      suppressible
      brat
      (Popkova et al., 2012)
      klu
      suppressible
      brat
      (Xiao et al., 2012)
      Nsf2
      suppressible
      brat
      (Laviolette et al., 2005)
      orb2
      suppressible
      brat
      (Santana and Casas-Tintó, 2017)
      Smox
      suppressible
      brat
      (Hevia et al., 2017)
      External Data
      Subunit Structure (UniProtKB)
      Interacts with nos and pum (PubMed:11274060). Acts via the formation of a quaternary complex composed of pum, nos, brat and the 3'-UTR mRNA of hb (PubMed:11274060). Not recruited by nos and pum to cyclin B 3'-UTR mRNA (PubMed:11274060). Might interact with mira; the interaction seems to be important for brat localization during mitosis (PubMed:16564014).
      (UniProt, Q8MQJ9 )
      Linkouts
      BioGRID - A database of protein and genetic interactions.
      DroID - A comprehensive database of gene and protein interactions.
      InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
      MIST (genetic) - An integrated Molecular Interaction Database
      MIST (protein-protein) - An integrated Molecular Interaction Database
      Pathways
      Signaling Pathways (FlyBase)
      Metabolic Pathways
      External Data
      Linkouts
      Genomic Location and Detailed Mapping Data
      Chromosome (arm)
      2L
      Recombination map

      2-54

      Cytogenetic map
      37C1-37C6
      Sequence location
      2L:19,133,816..19,179,758 [+]
      FlyBase Computed Cytological Location
      Cytogenetic map
      Evidence for location
      37C1-37C6
      Limits computationally determined from genome sequence between P{lacW}Ddck02104 and P{lacW}bratk06028&P{lacW}l(2)k09613k09613
      Experimentally Determined Cytological Location
      Cytogenetic map
      Notes
      References
      37C1-37D1
      (determined by in situ hybridisation)
      (Lukacsovich et al., 2001)
      37C1-37D1
      (Lukacsovich, 1999.1.13)
      37C6-37C7
      (determined by in situ hybridisation)
      (Spradling et al., 1999)
      37C5-37C6
      37C5--7 37C6--7
      (BDGP Project Members, 1994-1999)
      Experimentally Determined Recombination Data
      Location

      2-54

      (FlyBase, 2016)

      2-53.6

      (Comeron et al., 2012)
      Left of (cM)
      Right of (cM)
      Notes
      Stocks and Reagents
      Stocks (39)
      Genomic Clones (26)
      cDNA Clones (214)
       

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

      cDNA clones, fully 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
      GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
      Antibody Information
      Laboratory Generated Antibodies
       

      monoclonal

      (Shi et al., 2013)

      polyclonal

      (Shi et al., 2013)
      Commercially Available Antibodies
       
      Other Information
      Relationship to Other Genes
      Source for database identify of
      Source for database merge of

      Source for merge of: brat anon-37CDa

      (FlyBase, 1996-)
      Additional comments

      Source for merge of brat anon-37CDa was sequence comparison ( date:991108 ).

      (FlyBase, 1996-)
      Other Comments

      brat specifies intermediate neural progenitor identity through its B-box domains, downregulating the transcriptional activity of arm via a mechanism that is independent of asymmetric protein segregation.

      (Komori et al., 2014)

      brat regulates synapse development and endocytosis at the neuromuscular junction by suppressing translation of Mad.

      (Shi et al., 2013)

      Loss of brat in larvae causes neoplastic brain tumors. Fragments of brat brain tumours transplanted into adult hosts over-proliferate and kill their hosts within 2 weeks.

      (Beaucher et al., 2007)

      brat-induced metastases express both neuronal (elav) and glial cell (repo) type markers.

      (Beaucher et al., 2007)

      The brat gene product acts as a posttranscriptional inhibitor of dm in one of the two daughter cells of larval neuroblasts.

      (Betschinger et al., 2006)

      brat and 4EHP interact physically and both play a role in hb regulation.

      (Cho et al., 2006)

      S2 cells treated with dsRNA generated against this gene show reduced phagocytosis of Candida albicans compared to untreated cells.

      (Stroschein-Stevenson et al., 2006)

      brat appears to be involved in negatively regulating the level of cellular rRNA.

      (Frank et al., 2002)

      brat is a translational repressor.

      (Sonoda and Wharton, 2001)

      Located between coordinates +45.7 and +50.0.

      (Stathakis et al., 1995)

      Mutants display a brain neoplastic phenotype.

      (Mechler, 1994)

      Mutations at the brat locus cause defects in midoogenesis.

      (Schupbach and Wieschaus, 1991)

      Lethality occurs as pharate adults. Third-instar larvae of 13 of 14 alleles examined display enlarged brains -- up to eight times normal volume; greatest in presumptive optic center (Hankins and Wright). Brain fragments transplanted into normal adults show uncontrolled proliferation and metastatic invasion of other tissues (Gateff).

      Origin and Etymology
      Discoverer
      Etymology
      Identification
      External Crossreferences and Linkouts ( 105 )
      Sequence Crossreferences
      NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
      GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
      GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
      RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
      UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
      UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
      Other crossreferences
      BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
      Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
      Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
      Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
      Flygut - An atlas of the Drosophila adult midgut
      GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
      iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
      InterPro - A database of protein families, domains and functional sites
      KEGG Genes - Molecular building blocks of life in the genomic space.
      modMine - A data warehouse for the modENCODE project
      Linkouts
      BioGRID - A database of protein and genetic interactions.
      DroID - A comprehensive database of gene and protein interactions.
      DRSC - Results frm RNAi screens
      FLIGHT - Cell culture data for RNAi and other high-throughput technologies
      FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
      FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
      FlyMine - An integrated database for Drosophila genomics
      Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
      InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
      MIST (genetic) - An integrated Molecular Interaction Database
      MIST (protein-protein) - An integrated Molecular Interaction Database
      Synonyms and Secondary IDs (17)
      Reported As
      Symbol Synonym
      BRAT
      (Lefebvre and Lécuyer, 2018, Homem et al., 2015, Gonzalez, 2013, Kurusu et al., 2009)
      Brat
      (Liu et al., 2019, Malik et al., 2019, Piggott et al., 2019, Rivera et al., 2019, van den Ameele and Brand, 2019, Carmena, 2018, Olesnicky and Wright, 2018, Lou et al., 2017, Malik et al., 2017, Ma et al., 2016, Mukherjee et al., 2016, Yadav et al., 2016, Yasugi and Nishimura, 2016, Zhang et al., 2016, Bajaj et al., 2015, Jia et al., 2015, Schweisguth, 2015, Singh, 2015, Eroglu et al., 2014, Pargett et al., 2014, Bausek, 2013, Bergstralh et al., 2013, Gallant, 2013, Lu and Johnston, 2013, Homem and Knoblich, 2012, Kelsom and Lu, 2012, Marin, 2012, Saini and Reichert, 2012, Vazquez-Pianzola and Suter, 2012, Eliazer and Buszczak, 2011, Harris and Ashe, 2011, Miles et al., 2011, Weng and Lee, 2011, Hessle et al., 2009, Nie et al., 2009, Wang et al., 2009, Chia et al., 2008, Harris and Ashe, 2008, Sousa-Nunes et al., 2008, Speicher et al., 2008, Kadyrova et al., 2007, Neumüller et al., 2007, Slack et al., 2007, Speicher et al., 2007, Wang et al., 2007, Yamazaki et al., 2007, Betschinger et al., 2006, Bowman et al., 2006, Brun et al., 2006, Cho et al., 2006)
      CG10719
      (Yamaguchi and Takashima, 2018, Hevia et al., 2017, Christis and Munro, 2012, Müller et al., 2010, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Krejcí et al., 2009, Perkins et al., 2009, Berger et al., 2008, Sardiello et al., 2008, Quinones-Coello, 2007, Rehwinkel et al., 2006)
      Cf
      (McCrady and Tolin, 1994)
      E60
      anon-37CDa
      brat
      (Cho et al., 2020, Eichler et al., 2020, Guangming et al., 2020, Rastegari et al., 2020, Wei et al., 2020, Zraly et al., 2020, Chen and Read, 2019, Drummond-Barbosa, 2019, King et al., 2019, Luhur et al., 2019, Meltzer et al., 2019, Mirkovic et al., 2019, Mirzoyan et al., 2019, Molnar et al., 2019, Raza et al., 2019, Arbeille and Bashaw, 2018, Chai et al., 2018, Flora et al., 2018, Kang et al., 2018, Landskron et al., 2018, Loewen et al., 2018, Mora et al., 2018, Reichardt et al., 2018, Rossi et al., 2018, Wissel et al., 2018, Yamaguchi and Takashima, 2018, Banerjee and Roy, 2017, Janssens et al., 2017, Liu et al., 2017, Ma et al., 2017, Perry et al., 2017, Transgenic RNAi Project members, 2017-, Vallejos Baier et al., 2017, Wu et al., 2017, Narbonne-Reveau et al., 2016, Shlyueva et al., 2016, Bivik et al., 2015, Fernando et al., 2015, Gene Disruption Project members, 2015-, Gogendeau et al., 2015, Huang et al., 2015, Laver et al., 2015, Liu et al., 2015, Loedige et al., 2015, model organism Encyclopedia of Regulatory Network (modERN) Project, 2015-, Newton et al., 2015, Oktaba et al., 2015, Sinkovics, 2015, Weng and Cohen, 2015, Chen et al., 2014, Ferreira et al., 2014, Homem et al., 2014, Janssens et al., 2014, Komori et al., 2014, Komori et al., 2014, Luck et al., 2014, Miles et al., 2014, Olesnicky et al., 2014, Southall et al., 2014, Tipping and Perrimon, 2014, Xu et al., 2014, Chai et al., 2013, Jüschke et al., 2013, Neumüller et al., 2013, Shi et al., 2013, Southall et al., 2013, Adamson and Lajeunesse, 2012, Berger et al., 2012, Bunt et al., 2012, Carney et al., 2012, Hilgers et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Justiniano et al., 2012, Li et al., 2012, Mihailovich et al., 2012, Miles et al., 2012, Olesnicky et al., 2012, Popkova et al., 2012, Saini and Reichert, 2012, Smibert et al., 2012, Xiao et al., 2012, Abruzzi et al., 2011, Boyan and Reichert, 2011, Harris et al., 2011, Hilgers et al., 2011, Miles et al., 2011, Neumüller et al., 2011, Zhu et al., 2011, Chang et al., 2010, Janic et al., 2010, Müller et al., 2010, Rigoutsos, 2010, Temme et al., 2010, Ayroles et al., 2009, Flaherty et al., 2009, Krejcí et al., 2009, Robine et al., 2009, Saito et al., 2009, Sousa-Nunes et al., 2009, Stone and Ayroles, 2009, Basto et al., 2008, Berger et al., 2008, Bowman et al., 2008, FlyBase Curators, 2008-, Muraro et al., 2008, Neumuller et al., 2008, O'Farrell et al., 2008, Beaucher et al., 2007, Beaucher et al., 2007, Quinones-Coello, 2007, Tadros et al., 2007, Tountas and Fortini, 2007, Zhang et al., 2007, Bowler et al., 2006, Lee et al., 2006, Lee et al., 2006, Stroschein-Stevenson et al., 2006, Wang et al., 2006, Wodarz and Gonzalez, 2006, Beckstead et al., 2005, Caussinus and Gonzalez, 2005, Laviolette et al., 2005, Loop et al., 2004, McCrady and Tolin, 1994)
      fs(2)ltoPM43
      l(2)37Cf
      (Callahan et al., 1996, Callahan et al., 1995, Mechler, 1990, Gateff and Mechler, 1989, Wright, 1987, Pentz et al., 1986, Gilbert et al., 1984, Brittnacher and Ganetzky, 1983, Wright et al., 1981)
      l(2)E60
      (Wright et al., 1976)
      l(2)brat
      lincRNA.S1565
      (FlyBase Genome Annotators, 2014, Young et al., 2012)
      Name Synonyms
      Brain Tumor
      (Liu and Grosshans, 2017, Mukherjee et al., 2016, Eliazer and Buszczak, 2011, Lin, 2008, Speicher et al., 2008, Sonoda and Wharton, 2001)
      Brain tumor
      (Nguyen et al., 2016, Chen et al., 2014, Bausek, 2013, Vazquez-Pianzola and Suter, 2012, Eliazer and Buszczak, 2011, Krejcí et al., 2009, Chia et al., 2008, Beaucher et al., 2007, Cho et al., 2006, Lee et al., 2006)
      Brain tumour
      (Harris and Ashe, 2011)
      brain tumor
      (Mirzoyan et al., 2019, Huang et al., 2015, Southall et al., 2014, Justiniano et al., 2012, Miles et al., 2012, Saini and Reichert, 2012, Miles et al., 2011, Janic et al., 2010, Mastushita-Sakai et al., 2010, Temme et al., 2010, Flaherty et al., 2009, Robine et al., 2009, Stone and Ayroles, 2009, Basto et al., 2008, O'Farrell et al., 2008, Beaucher et al., 2007, Kankel et al., 2007, Tountas and Fortini, 2007, Bello et al., 2006, Brun et al., 2006, Wang et al., 2006, Beckstead et al., 2005, Loop et al., 2004)
      brain tumour
      (Southall and Brand, 2009)
      Secondary FlyBase IDs
      • FBgn0002034
      • FBgn0004827
      • FBgn0027355
      Datasets (0)
      Study focus (0)
      Experimental Role
      Project
      Project Type
      Title
      References (314)