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General Information
Symbol
Dmel\br
Species
D. melanogaster
Name
broad
Annotation Symbol
CG11491
Feature Type
FlyBase ID
FBgn0283451
Gene Model Status
Stock Availability
Gene Snapshot
In progress.Contributions welcome.
Also Known As
BR-C, Broad Complex, ecs, npr1, rbp
Key Links
Genomic Location
Cytogenetic map
Sequence location
X:1,575,132..1,664,869 [+]
Recombination map
1-0.2
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
Predictions / Assertions
-
Summaries
Gene Group (FlyBase)
C2H2 ZINC FINGER TRANSCRIPTION FACTORS -
Zinc finger C2H2 transcription factors are sequence-specific DNA binding proteins that regulate transcription. They possess DNA-binding domains that are formed from repeated Cys2His2 zinc finger motifs. (Adapted from PMID:1835093, FBrf0220103 and FBrf0155739).
Protein Function (UniProtKB)
Broad-complex proteins are required for puffing and transcription of salivary gland late genes during metamorphosis.
(UniProt, Q01295)
Broad-complex proteins are required for puffing and transcription of salivary gland late genes during metamorphosis.
(UniProt, Q24206)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
br: broad
thumb
br: broad
Edith M. Wallace, unpublished.
The br complementation group contains both amorphic and hypomorphic mutant alleles; amorphic alleles cause early prepupal developmental arrest; hypomorphic alleles cause late pupal or pharate adult developmental arrest or are viable. Null alleles display normal larval development but prevent elongation and eversion of discs giving rise to appendages in the pupal stage. Wings of the viable allele, br1, somewhat broader than normal; about 80% of normal length, with round full tip; crossveins closer together. Shape difference visible in middle prepupal stage immediately after eversion [Waddington, 1939, Proc. Nat. Acad. Sci. USA 25: 299-307; 1940, J. Genet. 41: 75-139 (fig.)]. A haplo-insufficient locus in that heterozygosity for a deficiency including the br locus leads to a slight br phenotype (Craymer and Roy, 1980, DIS 55: 200-04); furthermore, the deficiency in combination with br1 or br3 leads to drastic reduction in viability, especially at 18, and an extreme phenotype among survivors, including reduced palpi characteristic of rbp alleles, short rounded wings with interrupted veins, and malformed third legs, i.e., shortened and thickened femora and tibiae as well as misshapen basitarsi. The malformed-leg syndrome is enhanced by heterozygosity for Sb or sbd alleles (Beaton, Kiss, Fristrom, and Fristrom, 1988, Genetics 120: 453-64). br16/+ and Df(1)S39/+ display slight dominance of br effects in the presence of RpII215Ubl (Mortin and Lefevre, 1981, Chromosoma 82: 237-47).
l(1)2Bab
So named because of its failure to complement lethality of br and rbp mutations. Homozygotes and hemizygotes die in pupal stage; puparium formation delayed three [l(1)2Bab3] to six [l(1)2Bab1] hr. l(1)2Bab1 pupae have normal imaginal organs, and escapers have faded wings and reduced bristles on palpi. Prepupal lethal in combination with npr11 or Df(1)S39.
l(1)2Bc
Die in prepupal or early pupal stage after formation of a gas bubble. Imaginal discs fail to fuse, especially dorsally, to produce a continuous integument. Puparium formation variably delayed: 6 hr in l(1)2Bc1, 12 hr in l(1)2Bc2, and 9 hr in l(1)2Bc3 and l(1)2Bc4. Many late ecdysone puffs both in larvae and prepupae either absent or underdeveloped [l(1)2Bc1] [Zhimulev, Belyaeva, and Aizenzon, 1980, Genetika (Moscow) 16: 1613-31]. l(1)2Bc1 an amorphic allele fully complements the amorphic br5 (Kiss et al.).
l(1)2Bd
Males and homozygous females display normal phenotype and viability; however, in heterozygous combination with deficiencies or npr mutations, l(1)2Bd acts as a temperature-sensitive lethal; completely lethal at 29; at 25 or 18 most individuals die in late pupal stage, and survivors have faded wings, swollen abdomen, and reduced bristle number on palpi.
npr1: nonpupariating
Hemizygous male larvae fail to pupariate, although they survive 10-15 days after their normal sibs have pupariated. Four-day-old larvae appear normal as do their imaginal discs; normal ecdysteroid levels achieved. Discs become abnormal beginning on the sixth day; peripodial membrane becomes enormously distended and highly distorted; partially evaginated structure becomes visible in the disc lumen; do not undergo detailed morphological changes characteristic or metamorphosis, either in situ or in transplants into normal larvae [Fristrom, Fekete, and Fristrom, Wilhelm Roux's Arch. Dev. Biol. 190: 11-21 (fig.)]. Both salivary glands and fat bodies fail to undergo histolysis in situ or in vitro. Mutant flies able to produce ecdysone, but tissues unable to respond normally. In gynandromorphs, the female tissue forms a puparium, whereas npr1 male tissue remains larval; no adults survive (Kiss, Szabad, and Major, 1978, Mol. Gen. Genet. 164: 77-83; Kiss, Bencze, Fodor, Szabad, and Fristrom, 1975, Nature 262: 136-38). Implantation of wild-type ring glands into npr1 larvae does not rescue pupariation; however implanted wild-type or npr1 ring glands are able to rescue npr3 larvae [Kiss, Szabad, Belyaeva, Zhimulev, and Major, 1980, Development and Neurobiology of Drosophila (Siddiqi, Babu, Hall, and Hall, eds.) Plenum Press, New York and London, pp. 163-81]. No maternal effect of either npr13 or npr14 (Perrimon, Engstrom, and Mahowald, 1984, Dev. Biol. 105: 404-14). npr16 homozygous and hemizygous larvae die without exhibiting any sign of ecdysone-inducible puff formation; culture of slivary glands in 20-OH ecdysone produces partial development of some early, but none of late ecdysone-inducible puffs, and extraneous puff appears at 75CD (Belyaeva, Vlassova, Biyasheva, Kakpakov, Richards, and Zhimulev, 1981, Chromosoma 84: 207-19). npr1+ gene product also required for regression of the intermolt 68C glue puff [Belyaeva, et al. (npr6); Crowley, Mathers and Meyerowitz, 1984, Cell 39: 149-56 (npr3)] and for the transcription of the three 68C glue protein genes (Crowley et al).
rbp: reduced bristles on palpus
Late pupal lethal; most animals reach the pharate adult stage. rbp2 homozygotes and males survive with fewer than normal bristles on the palpus; when raised at 29, females exhibit faded wings and swollen abdomens; when raised at 18 males have faded wings. rbp2/Df(1)RA19 females virtually lethal; females carrying rbp2 and any of the other rbp alleles are completely viable when reared at 18, but at higher temperatures most die and escapers have reduced bristles on the palpus, shortened bristles on the scutellum, shrivelled or swollen abdomen shrivelled wings and eyes with crumpled surface.
Summary (Interactive Fly)
Gene Model and Products
Number of Transcripts
14
Number of Unique Polypeptides
6

Please see the GBrowse view of Dmel\br or the JBrowse view of Dmel\br 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
Gene model reviewed during 6.06
Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated
Gene model reviewed during 5.44
gene_with_stop_codon_read_through ; SO:0000697
Gene model reviewed during 5.39
Gene model reviewed during 5.42
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.45
Stop-codon suppression (UAA) postulated; FBrf0216884.
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0070265
8223
702
FBtr0070263
7839
880
FBtr0070261
3867
880
FBtr0070262
4419
724
FBtr0070266
4236
663
FBtr0300427
4305
880
FBtr0300428
3981
880
FBtr0300429
3119
514
FBtr0303562
9822
514
FBtr0303563
4213
724
FBtr0303564
4546
724
FBtr0308319
4368
663
FBtr0330406
4188
1011
FBtr0332293
4235
880
Additional Transcript Data and Comments
Reported size (kB)
11.5, 9.5, 7.4, 6.5, 4.5, 2 (northern blot)
5.2 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0070255
74.1
702
7.70
FBpp0070253
92.3
880
7.98
FBpp0070251
92.3
880
7.98
FBpp0070252
77.1
724
7.87
FBpp0070256
70.5
663
7.49
FBpp0289656
92.3
880
7.98
FBpp0289657
92.3
880
7.98
FBpp0289658
55.5
514
7.12
FBpp0292595
55.5
514
7.12
FBpp0292596
77.1
724
7.87
FBpp0292597
77.1
724
7.87
FBpp0300638
70.5
663
7.49
FBpp0303432
105.5
1011
7.57
FBpp0304572
92.3
880
7.98
Polypeptides with Identical Sequences

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

880 aa isoforms: br-PB, br-PC, br-PH, br-PI, br-PQ
724 aa isoforms: br-PE, br-PM, br-PN
663 aa isoforms: br-PG, br-PO
514 aa isoforms: br-PJ, br-PL
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Crossreferences
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\br using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (25 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Biological Process (22 terms)
Terms Based on Experimental Evidence (18 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from expression pattern
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:twi; FB:FBgn0003900
inferred from genetic interaction with FLYBASE:Mef2; FB:FBgn0011656
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN001135010
(assigned by GO_Central )
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
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: reference states 4 hours before puparium formation

radioisotope in situ
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Under conditions of nutritional shortage, initial expression detected later, at stage 6. Expression of different isoforms varies; BR-C Z2 and BR-C Z3 appear to mediate response to varying nutritional levels.
br transcript is localized to neuronal cell bodies in all parts of the adult brain. Expression levels in the adult optic lobe are lower than in the brain and subesophageal ganglion, with a region of particular intensity in a cluster of cells at the top of the cervical connective as it joins the subesophageal ganglion, and in cells surrounding the calyces of the mushroom bodies.
RNA blots were carried out on RNA extracted from staged larval and prepupal salivary glands. br transcripts are induced in apparent response to the late-larval and prepupal ecdysone pulses paralleling the puffing response at 2B5. This response is enhanced in response to ectopic ftz-f1 expression.
The major 4.5kb and minor 2, 7.4, 9.5, and 11.5kb br transcripts are present in ovaries and at low levels in embryonic and early larval stages. They rise dramatically in third instar through prepupal stages, consistent with being ecdysone-inducible.
Probes which hybridize to the 4.5kb, 7.4kb and 4.5kb br transcripts in the parent strain, detect transcripts of 6.5kb, 6.5kb and 5.2kb, respectively in mutant brnpr-fs ovaries.
Probes which hybridizes to the 4.5kb, 7.4kb and 4.5kb br transcripts in the parent strain, detect transcripts of 6.5kb, 6.5kb and 5.2kb, respectively in brnpr-fs ovaries.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
distribution deduced from reporter
Stage
Tissue/Position (including subcellular localization)
Reference
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
neuron | subset of prothoracic neuromere

Comment: Assay specific to Z3 isoform (br-RA). 72h-120h AEL

neuron | subset of metathoracic neuromere

Comment: Assay specific to Z3 isoform (br-RA). 72h-120h AEL

neuron | subset of mesothoracic neuromere

Comment: Assay specific to Z3 isoform (br-RA). 72h-120h AEL

neuron | subset of lamina

Comment: Assay specific to Z3 isoform (br-RA). Until 14h APF.

neuron | subset of medulla

Comment: Assay specific to Z3 isoform (br-RA). Until 27h APF.

salivary gland

Comment: reference states 0-6 hr APF

neuron | subset of embryonic/larval central nervous system

Comment: Assay specific to Z3 isoform (br-RA). 5.5h-16h APF.

glial cell | subset of ventral nerve cord

Comment: Assay specific to Z4 isoforms (br-RB,RC,RH,RI,RP,RQ). 5.5h-94h APF.

neuron | subset of ventral nerve cord

Comment: Assay specific to Z4 isoforms (br-RB,RC,RH,RI,RP,RQ). 5.5h-94h APF.

neuron | subset of lobula complex

Comment: Assay specific to Z3 isoform (br-RA). 27h APF.

neuron | subset of embryonic/larval mushroom body

Comment: Assay specific to Z3 isoform (br-RA). 95h APF.

western blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
br protein is detected in the embryonic central nervous system and in a subset of neurons of the thoracic and abdominal neuromeres just after hatching. Specific labelling for the Z3 isoform (br-RA) shows expression in neuroblast clones in the thoracic neuromeres: at the start of late third instar (72h AEL) arrested embryonic-born secondary neurons strongly express Z3 isoform in the oldest neurons of the cluster (basal) whereas expression is absent from the younger neurons (apical). Weak expression of Z3 isoform in larval-born secondary neurons is observed in the oldest neurons of the cluster at 72h AEL, becoming stronger as late third instar progresses, and disappears by pupariation. At this stage, Z3 isoform is observed in the younger neurons of cluster, except for those nearest to the neuroblast. In late larval and pupal stages, Z3 isoform is detected in the optic lobe and in four neuron clusters in the mushroom bodies. Specific labelling for the Z1 isoforms (br-RE,RG,RM,RN,RO) or Z4 isoforms (br-RB,RC,RH,RI,RP,RQ) is observed in neurons and glia in the central nervous system in pupal stages. The neurons labelled by Z3 isoform antibody correspond to the neurons that are labelled by the antibody for br protein; the subset of glia and neurons marked by Z1 or Z4 isoforms correspond to smaller subsets of the br positive cells.
An antibody directed to the core domain shared by all of the protein isoforms labeled over 300 distinct areas on larval and prepupal salivary glands. Labeling was observed in large and small puff regions as well as interbands. Pericentric heterochromatic labeling was observed only in one region - 20D and 20E of the X chromosome. This staining correlates with that observed using an antibody specific for the Z1 isoform which has been reported to be the predominate br isoform in larval salivary glands. In contrast, Z2 and Z3 specific antibodies did not label salivary gland polytene chromosomes.
Expression of br protein in larval prothoracic gland is isoform-specific. The Z2 and Z3 isoforms are detectable 17 hours after ecdysis into the third instar (89 hr AEL), and continue to be expressed through the white prepupa stage. The Z4 isoform is intensely expressed in wandering third instar larvae and white prepupae; and the Z1 isoform initiates intense expression later in the third instar.
At wandering third instar larval stage, br protein is widely expressed in the central nervous system. Staining localized to the nuclei of central nervous system, with prominent labeling in the optic lobes, brain, subesophageal ganglion, and thoracic segments of the ventral nerve cord. Expression is also observed in cells at the midline of the ventral nerve cord. Staining is absent in the neuropil.
Marker for
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
Reporter: P{br-GAL4.CR}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GawB}brPG12
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\br in GBrowse 2
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
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
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Images
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 124 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 26 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of br
Transgenic constructs containing regulatory region of br
Deletions and Duplications ( 65 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal (with br6)
lethal (with br7)
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
maxillary palpus & macrochaeta
microchaeta & adult thorax
microchaeta & thorax
muscle tendon junction & coxal tergal remotor muscle
muscle tendon junction & lateral oblique dorsal muscle
muscle tendon junction & tergosternal muscle
ovary & nucleus | conditional cs
sensory neuron & dorsal mesothoracic disc, with Scer\GAL4bbg-C96
wing (with br7)
wing margin & sensory neuron, with Scer\GAL4bbg-C96
wing margin & sensory neuron | somatic clone
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (6)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
No
1 of 15
Yes
Yes
1 of 15
Yes
No
 
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
Yes
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (0)
No records found.
Rattus norvegicus (Norway rat) (0)
No records found.
Xenopus tropicalis (Western clawed frog) (0)
No records found.
Danio rerio (Zebrafish) (1)
1 of 15
Yes
No
Caenorhabditis elegans (Nematode, roundworm) (0)
No records found.
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG0919069G )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila simulans
Drosophila erecta
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091506O3 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W09XF )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis florea
Little honeybee
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Apis mellifera
Western honey bee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Bombus terrestris
Buff-tailed bumblebee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X09TA )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
No non-Arthropod Metazoa orthologies identified
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (22)
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
2 of 10
2 of 10
2 of 10
2 of 10
1 of 10
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    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
    Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
     
    Disease Associations of Human Orthologs (via DIOPT v7.1 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
    esyN Network Diagram
    Show neighbor-neighbor interactions:
    Select Layout:
    Legend:
    Protein
    RNA
    Selected Interactor(s)
    Interactions Browser

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    RNA-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    esyN Network Diagram
    esyN Network Key:
    Suppression
    Enhancement

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    External Data
    Linkouts
    BioGRID - A database of protein and genetic 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
    Gene Group - Pathway Membership (FlyBase)
    External Data
    Linkouts
    KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    X
    Recombination map
    1-0.2
    Cytogenetic map
    Sequence location
    X:1,575,132..1,664,869 [+]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    2B3-2B4
    Limits computationally determined from genome sequence between P{EP}brEP1515 and P{EP}EP1444&P{EP}CG14818EP1190
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    2B1-2B8
    (determined by in situ hybridisation) 2B1--4 (determined by in situ hybridisation)
    Location based on deficiency mapping.
    2B1-2B8
    (determined by in situ hybridisation)
    2B1-2B4
    (determined by in situ hybridisation)
    2B5-2B5
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Notes
    Recombinational mapping experiments show that the 'rbp', 'l(1)2Bc' and 'broad' class alleles are separable by recombination.
    Mapping based on 15 crossovers between brnpr-4 and y, 36 between brnpr-4 and w and 159 between brnpr-4 and ec.
    brnpr-3 maps to 1-0.2, brnpr-4 maps to 1-0.4.
    Stocks and Reagents
    Stocks (59)
    Genomic Clones (24)
    cDNA Clones (57)
     

    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 sequences
    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)
    RNAi and Array Information
    Linkouts
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    Antibody Information
    Laboratory Generated Antibodies
    Commercially Available Antibodies
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for database merge of
    Source for merge of: br CG11491
    Source for merge of: br CG11514
    Source for merge of: br l(1)G0318 l(1)G0401 l(1)G0018 l(1)G0042 l(1)G0284a
    Source for merge of: br CG11511
    Source for merge of: br CG11509
    Additional comments
    Annotation CG11491 merged with CG11509 in release 6.06 of the genome annotation; CG11509 corresponds to a portion of the 3' UTR of an alternative transcript isoform of br.
    Annotations CG11491, CG11514 and CG11511 merged as CG11491 in release 3 of the genome annotation.
    Russian investigators originally defined four mutually complementing, lethally mutable loci, which function in ecdysone-dependent induction of metamorphosis: br, broad, rbp, reduced bristles on palpus, l(1)2Bc, and l(1)2Bd. Based on amorphic mutations, defined on the basis of having equivalent phenotypes in homozygous and hemizygous females, Kiss et al. (FBrf0048202) define two complementation groups: br, comprising br, rbp and l(1)2Bd (Belyaeva et al., FBrf0045819), and l(1)2Bc; npr alleles are non-complementing.
    Other Comments
    Annotation (CG11509) restored in release 5.33 of the genome annotation.
    Annotation (CG11509) eliminated in release 5.2 of the genome annotation.
    The br product prevents rho expression in egg chambers.
    br appears to be required for Ddc activation eclosion.
    The ecdysone pathway is required for proliferation and differentiation of hematopoitic precurssors of lamellocytes and crystal cells.
    br isoforms regulate morphogenetic furrow progression and photoreceptor specification in the developing eye.
    br is required for salivary gland, autophagic, cell death.
    rbp+ function is required in the dorsal epidermal muscle attachment site for the correct attachment of the indirect flight muscles.
    The target genes of the BR-C (br) in oogenesis include a protein essential for endoreplication and chorion gene amplification.
    br encodes a family of zinc-finger DNA binding proteins generated by alternative splicing, which produces four classes of proteins; Z1, Z2, Z3 and Z4. The Z1 isoform provides full rbp+ function. Z2 provides br+ function (Z2 is the only protein isoform to provide br+ function), and partial 2Bc+ function. Z3 provides full 2Bc+ function. Z4 provides partial rbp+ function.
    br is expressed in a bilaterally symmetrical pattern in the lateral-dorsal-anterior follicle cells during late oogenesis.
    br is essential for the morphogenesis of dorsal appendages.
    Wild-type rbp function is required for the correct and stable attachment of the thoracic muscles to the body wall, fasciculation and maintenance of these thoracic muscles. The primary effect of rbp mutations is to disrupt muscle attachment, causing muscles to choose incorrect sites or to make weak attachments that ultimately fail.
    The ecdysone response of individual br zinc finger RNA isoforms is examined in cultured imaginal discs. Studies reveal a complex temporal pattern of RNA expression.
    The br complex (BR-C) early gene encoded by the Z1 isoform (rbp) directly activates late gene transcription (genes from the 71E salivary glands late puff) by interacting with late gene cis-acting regulatory elements and this interaction is responsible for the temporal linkage of early and late ecdysone-induced gene expression.
    br gene participates in parallel tissue-specific ecdysone response cascades to Hsp23.
    A transgene containing the Z1 isoform can rescue a mutant 'rbp' phenotype. A transgene containing the Z2 isoform can rescue a mutant br phenotype and can partially rescue the mutant lethality of 'l(1)2Bc'. A transgene containing the Z3 isoform can rescue the mutant lethality of 'l(1)2Bc'.
    P-element rescue constructs suggest that the Z1 isoform of the BRC corresponds to 'rbp' function, and the Z2 isoform to 'br' function, but that there is no simple correlation between any one BRC protein and 'l(1)2Bc' function.
    Analysis of morphological and molecular phenotypes of double mutants between alleles of br and Eip74EF reveals that br and Eip74EF share functions in puparium formation, pupation and early gene induction. The br and Eip74EF transcription factors may directly interact to regulate the expression of salivary gland glue and late genes.
    Hormonal induction of Ddc in the epidermis is mediated by br.
    ImpE1 and Dfd have been examined for their positions relative to the Broad complex genes in the hormone-regulated pathway of CNS metamorphosis. Activity of any individual Broad complex subfunction is not required for ImpE1 induction.
    Mapping of DNase hypersensitive sites (DHS) in mutants reveals changes in chromatin structure associated with sites that presumably contain target sequence for the BR-C gene products.
    Differential expression of Broad complex transcription factors may forecast tissue-specific developmental fates during metamorphosis.
    br function is required for the proper timing of Sgs4 induction and acts through sequences located outside the Sgs4-Pig1 intergenic region.
    Ecdysteroid-regulated gene.
    Both 'rbp' and 'l(1)2Bc' are required for glue gene induction in mid-third instar larvae. The intermolt secondary-response genes and the late secondary-response genes are absolutely dependent on br for the induction. In addition the 'l(1)2Bc' function is required for glue gene repression in prepupae and the complete ecdysone induction of early mRNAs from Eip74EF, Eip75B and the broad complex itself.
    Mosaic experiments reveal a somatic but not germ line requirement for npr function.
    Developmentally regulated peak of arginine kinase at pupal stage P3 is affected by altering copy number of Eip75B and BRC, but not Eip74EF.
    The temporal pattern of br transcription during the third larval instar stage has been analysed.
    Localisation of hfw, dor and br are clustered in the X chromosomes of taxonomically distant Drosophila species, suggesting an evolutionary significance of such an organisation.
    Mutations at br disrupt the metamorphosis of internal tissues.
    When the 2B region is subjected to position effect the br locus in inactivated.
    The mlf phenotype (malformed syndrome), wing malformations and leg defects, can be produced by a dominant genetic interaction between Sb and br.
    'rbp' function is necessary for intermoult and late transcription in salivary glands. Transcription of genes in the ecdysone-induced puffing cascade is dependent on 'rbp' function.
    The npr wild type gene product, which is necessary for the activation of intermoult glue genes, is necessary for the inactivation of the pre-moult genes.
    br has been located within a puff on the telomeric chromosome of D.funebris, D.virilis, D.hydei, D.repleta, D.mercatorum and D.paranaensis, within a puff on the distal part of the X chromosome of D.kanekoi and within a puff on the proximal portion of the X chromosome of D.pseudoobscura.pseudoobscura.
    Ovaries from fertile females transplanted into br mutant females succeeded in connecting to recipient oviducts suggesting that female sterile mutations at the br locus are somatic line specific: abnormal morphogenesis of their genital disc is due to loss of normal sensitivity to ecdysterone.
    The steroid hormone 20-hydroxyecdysone can regulate RNA levels of a single gene both positively and negatively depending on hormone concentration. br can modify the direction of the gene's response to a hormone signal.
    Mutations at br reduce the transcription rate or stability of the small heat shock protein mRNAs.
    Female sterile mutations at the br locus do not interfere with the expression of Cp70.
    'l(1)2Bc' is involved in imaginal disc fusion. br is involved in imaginal disc evagination.
    'br' and 'l(1)2Bc' alleles are fully complementing.
    'l(1)2Bc' function is involved in appendage elongation.
    The br complementation group contains both amorphic and hypomorphic mutant alleles; amorphic alleles cause early prepupal developmental arrest; hypomorphic alleles cause late pupal or pharate adult developmental arrest or are viable. Null alleles display normal larval development but prevent elongation and eversion of discs giving rise to appendages in the pupal stage. Wings of the viable allele, br1, somewhat broader than normal; about 80% of normal length, with round full tip; crossveins closer together. Shape difference visible in middle prepupal stage immediately after eversion (FBrf0004792; FBrf0005070). A haplo-insufficient locus in that heterozygosity for a deficiency including the br locus leads to a slight br phenotype (FBrf0034402); furthermore, the deficiency in combination with br1 or br3 leads to drastic reduction in viability, especially at 18oC, and an extreme phenotype among survivors, including reduced palpi characteristic of 'rbp' alleles, short rounded wings with interrupted veins and malformed third legs, i.e., shortened and thickened femora and tibiae as well as misshapen basitarsi. The malformed-leg syndrome is enhanced by heterozygosity for Sb alleles (FBrf0048216). br16/+ and Df(1)S39/+ display slight dominance of br effects in the presence of RpII215Ubl (FBrf0035975). Hemizygous 'npr' male larvae fail to pupariate, although they survive 10-15 days after their normal sibs have pupariated. Four-day-old larvae appear normal as do their imaginal discs; normal ecdysteroid levels achieved. Discs become abnormal beginning on the sixth day; peripodial membrane becomes enormously distended and highly distorted; partially evaginated structure becomes visible in the disc lumen; do not undergo detailed morphological changes characteristic or metamorphosis, either in situ or in transplants into normal larvae (FBrf0037303). Both salivary glands and fat bodies fail to undergo histolysis in situ or in vitro. 'npr' mutant flies able to produce ecdysone, but tissues are unable to respond normally. In gynandromorphs, the female tissue forms a puparium, whereas 'npr' male tissue remains larval; no adults survive (FBrf0032169; FBrf0029263). Implantation of wild-type ring glands into 'npr' larvae does not rescue pupariation; however implanted wild-type or 'npr' ring glands are able to rescue brnpr-3 larvae (FBrf0034101). 'npr' alleles fail to show regression of the intermolt 68C glue puff (FBrf0035971; FBrf0040472) or induction of ecdysone induced 'early' puffs in larval salivary gland polytene chromosomes. 'rbp' alleles are late pupal lethal; most animals reach the pharate adult stage. Viable 'rbp' adults have a reduced number of bristles on the maxillary palpus, and may show other bristle and wing defects.
    br encodes a family of Zinc-finger proteins that possess both common and unique exons. The gene is expressed as an 'early' ecdysone puff in 2B1--2B5 in salivary gland polytene chromosomes. A consequence of the molecular organization of the br gene is a complex pattern of complementation between mutant alleles.
    Origin and Etymology
    Discoverer
    Etymology
    Identification
    External Crossreferences and Linkouts ( 148 )
    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 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.
    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
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    KEGG Genes - Molecular building blocks of life in the genomic space.
    KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
    Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Synonyms and Secondary IDs (108)
    Reported As
    Symbol Synonym
    BR-C
    (Lv et al., 2019, Lee et al., 2018, Velentzas et al., 2018, Yun et al., 2017, Chaharbakhshi and Jemc, 2016, Sengupta et al., 2016, Nicolson et al., 2015, Mulakkal et al., 2014, Deng and Kerppola, 2013, Denton et al., 2013, Denton et al., 2013, Hyun, 2013, McDermott and Davis, 2013, Anhezini et al., 2012, Ihry et al., 2012, Jin et al., 2012, Avery et al., 2011, Farkaš et al., 2011, Johnston et al., 2011, Lin et al., 2011, Miles et al., 2011, Stepchenko et al., 2011, Dialynas et al., 2010, Francis et al., 2010, Rendina et al., 2010, Ritter and Beckstead, 2010, Sawatsubashi et al., 2010, Bernardo et al., 2009, Chittaranjan et al., 2009, Kozlova et al., 2009, Lachance et al., 2009, Cakouros et al., 2008, Chen et al., 2008, Hoopfer et al., 2008, Jindra and Konopova, 2008, Thomas and Cripps, 2008, Bashirullah et al., 2007, Cao et al., 2007, McBrayer et al., 2007, Peretz et al., 2007, Shravage et al., 2007, Thummel, 2007, Atkey et al., 2006, Choi et al., 2006, Ciurciu et al., 2006, Farkas et al., 2006, Furuhashi et al., 2006, Kelly and Daniel, 2006, Legube et al., 2006, Badenhorst et al., 2005, Beckstead et al., 2005, Caceres and Nilson, 2005, Farkas et al., 2005, Kilpatrick et al., 2005, Dubrovsky et al., 2004, Martin and Baehrecke, 2004, Mazo et al., 2004, Terashima and Bownes, 2004, Chiang and Kurnit, 2003, Daish et al., 2003, James and Berg, 2003, Kumar et al., 2003, Nakamura and Matsuno, 2003, Berger et al., 2002, Bialecki et al., 2002, Cakouros et al., 2002, Duncan and Warrior, 2002, Gonzalez et al., 2002, Gonzy et al., 2002, Lee et al., 2002, Lee et al., 2002, Pokholkova and Zhimulev, 2002, Thummel and Chory, 2002, Zhang et al., 2002, Zhou and Riffiford, 2002, Beckstead et al., 2001, Deng et al., 2001, Dubrovsky et al., 2001, Farkas and Mechler, 2001, Lee and Baehrecke, 2001, Lee et al., 2001, Lee et al., 2001, Simon et al., 2001, Suzanne et al., 2001, Thummel, 2001, Zhou and Riddiford, 2001, Baehrecke, 2000, Buszczak and Segraves, 2000, Hsu and Schulz, 2000, Jiang et al., 2000, Kozlova and Thummel, 2000, Lafont, 2000, Lee and Baehrecke, 2000, Lee et al., 2000, Li and Bender, 2000, Peri and Roth, 2000, Yamada et al., 2000, Baehrecke and Lee, 1999, Broadus et al., 1999, Buszczak et al., 1999, Lam et al., 1999, Roth et al., 1999, Roy and VijayRaghavan, 1999, Zhao and Bownes, 1999, Brennan et al., 1998, Buszczak and Segraves, 1998, D'Avino and Thummel, 1998, Deng and Bownes, 1998, Doronkin and Korochkin, 1998, Fisk and Thummel, 1998, Hall and Thummel, 1998, Bayer et al., 1997, Bender et al., 1997, Lam et al., 1997, Thummel, 1997, Baehrecke, 1996, Bayer, 1996.3.16, Ito et al., 1996, Makunin et al., 1996, D'Avino et al., 1995, Atkinson, 1994, Huet et al., 1993, Karim and Thummel, 1992, Restifo et al., 1992, Galceran et al., 1990)
    EG:123F11.1
    br
    (Jeong et al., 2019, Shih et al., 2019, Shokri et al., 2019, Davie et al., 2018, Hu et al., 2017.6.13, Kang et al., 2017, Osterfield et al., 2017, Revaitis et al., 2017, Djabrayan and Casanova, 2016, Guo et al., 2016, Jia et al., 2016, Kwon et al., 2016, Doggett et al., 2015, FlyBase Genome Annotators, 2015, Jia et al., 2015, Katsuyama et al., 2015, Nadimpalli et al., 2015, Schertel et al., 2015, Ugrankar et al., 2015, Xie et al., 2015, Boyle et al., 2014, Fauré et al., 2014, Haelterman et al., 2014, Huang et al., 2014, Ihry and Bashirullah, 2014, Jia et al., 2014, Liang et al., 2014, Luhur et al., 2014, Shlyueva et al., 2014, Slattery et al., 2014, Cheung et al., 2013, Enuameh et al., 2013, Fleming and McShea, 2013, Iyer et al., 2013, McKay and Lieb, 2013, Riddiford, 2013.12.3, Spokony and White, 2013.7.5, Vanden Broeck et al., 2013, Wang et al., 2013, Yamamoto et al., 2013-, Fuchs et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Niepielko et al., 2012, Simakov et al., 2012, Spokony and White, 2012.5.22, Stern et al., 2012, Abdou et al., 2011, Abdou et al., 2011, Jungreis et al., 2011, Seong et al., 2011, Weake et al., 2011, Zheng et al., 2011, Ables and Drummond-Barbosa, 2010, Kim et al., 2010, Xiang et al., 2010, Bai et al., 2009, Bernardi et al., 2009, Cui et al., 2009, Dalton et al., 2009, Dworkin et al., 2009, Lembong et al., 2009, Mirth et al., 2009, Parrish et al., 2009, Perkins et al., 2009.8.10, Spokony and Restifo, 2009, Zartman et al., 2009, Zhou et al., 2009, Christensen et al., 2008.12.28, Cook et al., 2008.9.3, Gilchrist et al., 2008, Khokhar et al., 2008, Lembong et al., 2008, Liu and Lehmann, 2008, Minakuchi et al., 2008, Pearse et al., 2008, Stofanko et al., 2008, Wang et al., 2008, Weake et al., 2008, Yakoby et al., 2008, Yakoby et al., 2008, Zartman et al., 2008, Zhang et al., 2008, Zhou and Riddiford, 2008, Aerts et al., 2007, Cao et al., 2007, Grieder et al., 2007, Zeitouni et al., 2007, Zhang et al., 2007, Zhou et al., 2007, Alekseyenko et al., 2006, Armstrong et al., 2006, Basso et al., 2006, Chen and Schupbach, 2006, Hammonds and Fristrom, 2006, Manak et al., 2006, Wilson et al., 2006, Consoulas et al., 2005, Zhou et al., 2004, Riddiford et al., 2003, Klebes et al., 2002, Sorrentino et al., 2002)
    l(1)2Ba
    l(1)2Bad
    l(1)2Bb
    l(1)G0284
    l(1)d norm-12
    l(1)d.norm.1
    l(1)dn1
    l(1)n34
    l(1)npr1
    l(1)ts132
    l(1)ts144
    l(1)ts358
    l(1)ts376
    o.c.c.
    rdp
    Name Synonyms
    ecdysone-sensitivity
    lethal(1)2Bab
    lethal(1)2Bc
    lethal(1)2Bd
    lethal(1)discs normal
    lethal(1)non-pupariating-1
    non-pupariating
    non-pupariating-1
    overlapping-complementation-complex
    reduced bristles on palps
    reduced palps
    unequal wings
    Secondary FlyBase IDs
    • FBgn0000210
    • FBgn0000801
    • FBgn0001373
    • FBgn0001374
    • FBgn0001375
    • FBgn0001644
    • FBgn0001684
    • FBgn0002964
    • FBgn0003222
    • FBgn0005389
    • FBgn0005581
    • FBgn0010011
    • FBgn0026711
    • FBgn0027219
    • FBgn0027234
    • FBgn0028290
    • FBgn0028364
    • FBgn0029583
    • FBgn0029585
    • FBgn0029586
    • FBgn0029584
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    References (667)