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Gene Dmel\br

General Information
Symbol	Dmel\br	Species	D. melanogaster
Name	broad	Annotation symbol	CG11491
Feature type	protein_coding_gene	FlyBase ID	FBgn0000210
Gene Model Status	Current	Stock availability	49 publicly available
Also Known As	BR-C, ecs, BRC, npr1, rbp, 2Bc, BrC-Z1, l(1)PP1, BR-C Z1, Z4
Genomic Location
Chromosome (arm)	X	Recombination map	1-0.28
Cytogenetic map	2B3-2B4	Sequence location	X:1,469,165..1,558,700 [+]
Genomic Maps Select View:Gene Models/EvidenceExpression/RegulationMegaViewGene Disruptions, Stocks and Reagents modENCODE GBrowse	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene broad is referred to in FlyBase by the symbol Dmel\br (CG11491, FBgn0000210). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: RNA polymerase II regulatory region sequence-specific DNA binding; DNA binding. There is experimental evidence for 15 unique biological process terms, many of which group under: anatomical structure development; biological regulation; response to hormone stimulus; sensory organ development; organ morphogenesis; multi-organism process; multicellular organismal reproductive process; catabolic process; eclosion; response to chemical stimulus; gland morphogenesis. 141 alleles are reported. The phenotypes of these alleles are annotated with: organ system subdivision; adult segment; adult mesothoracic segment; gland; organ system; egg; lobula complex; external compound sense organ; mouthpart; indirect flight muscle. It has 13 annotated transcripts and 13 annotated polypeptides. Protein features are: BTB/POZ; BTB/POZ fold; BTB/POZ-like; Zinc finger, C2H2; Zinc finger, C2H2-like. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of high expression to a trough of extremely low expression. Peak expression observed during late larval stages, during early pupal stages. Summary of FlyAtlas Anatomical Expression Data: Two or more Affy2 ProbeSets identify exons of this gene. This is a summary of the tissue expression peaks exhibited in at least one of these ProbeSets. Expression at high levels in the following post-embryonic organs or tissues: larval salivary gland. Expression at moderate levels in the following post-embryonic organs or tissues: larval/adult central nervous system, larval hindgut, larval Malpighian tubules, larval fat body, larval trachea, larval carcass. Gene sequence location is X:1469165..1558700.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	br: broad 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.
Recent Updates
Description	What does this section display? This section contains items that were added to this record for each release. It currently only tracks new links between this FlyBase report and other FlyBase data classes (e.g. genes, references, stocks) or controlled vocabulary terms (e.g. GO, anatomy terms). What does this section not display? This section does not currently display links that were removed or gene model changes.
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FB2011_10	References Herranz et al., 2010, Mol. Ecol. 19(19): 4255--4264
FB2012_01	Unknown western blot Sequence features HFA18743 HFA17789 HFA18744 BKN31704 HFA17691 Controlled Vocabulary Terms cellular response to ecdysone day 1 of adulthood prepupal stage P4(ii) wandering third instar larval stage third instar larval stage L1 negative regulation of gene expression third instar - uncleared gut stage adult abdomen embryonic/larval central nervous system neuronal cell body day 2 of adulthood adult thorax third instar - partially cleared gut stage third instar - cleared gut stage ventral nerve cord prepupal stage P4(i) third instar larval stage L2 late third instar larval stage embryonic/larval optic lobe pupal stage P5(i) positive regulation of transcription from RNA polymerase II promoter RNA polymerase II regulatory region sequence-specific DNA binding adult brain pupal stage P5(ii) prepupal stage P4 adult head References Gradilla et al., 2011, Genetics 189(3): 871--883 Zartman et al., 2011, Phys. Biol. 8(4): 045003
All updates	Click here to see a list of all updates to this record from FB2010_08 and on.
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 2B3-2B4
Experimentally Determined Cytological Location
Cytogenetic map Notes References 2B1-2B8   (determined by in situ hybridisation) 2B1--4 (determined by in situ hybridisation)   (Peter et al., 2002) 2B1-2B8   (determined by in situ hybridisation)   (Schaefer et al., 1999.8.12) 2B1-2B8   (determined by in situ hybridisation)   (Schaefer et al., 1999.6.15) 2B1-2B4   (determined by in situ hybridisation)   (Schaefer et al., 1999.3.29) 2B5-2B5   (determined by in situ hybridisation)   (DiBello et al., 1991) 2B3-2B5   Location based on deficiency mapping.   (Orr et al., 1989)
Experimentally Determined Recombination Data
Location	1-0.2--0.4 (Kiss et al., 1978) 1-0.2 (Gatti and Baker, 1989) 1- (Orr et al., 1989) 1-0.28   1-0.43 (Belyaeva et al., 1981) 1-0.5 +/- 0.2 (Fahmy and Fahmy, 1958) 2-104.5 (Buzzati-Traverso, 1947)
Left of (cM)	dor hfw (Aizenzon and Belyaeva, 1982) Pgd (Gvozdev et al., 1970) w (Belyaeva et al., 1981) ec (Orr et al., 1989)
Right of (cM)	y (Belyaeva et al., 1981) sc (Orr et al., 1989) sta (Aizenzon and Belyaeva, 1982)
Notes	brnpr-3 maps to 1-0.2, brnpr-4 maps to 1-0.4. (Kiss et al., 1978) Mapping based on 15 crossovers between brnpr-4 and y, 36 between brnpr-4 and w and 159 between brnpr-4 and ec. (Belyaeva et al., 1981) Recombinational mapping experiments show that the 'rbp', 'l(1)2Bc' and 'broad' class alleles are separable by recombination. (Aizenzon and Belyaeva, 1982)
Gene Model & Products
Please see the GBrowse view of Dmel\br for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
	Gene model reviewed during 5.39 Gene model reviewed during 5.42
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) br-RA FBtr0070265  NM_166894   8021   702 br-RB FBtr0070263  NM_166896   4155   880 br-RC FBtr0070261  NM_166897   3867   880 br-RE FBtr0070262  NM_080020   4419   724 br-RG FBtr0070266  NM_166898   4236   663 br-RH FBtr0300427  NM_001144672   4305   880 br-RI FBtr0300428  NM_166895   3981   880 br-RJ FBtr0300429  NM_166893   3119   514 br-RK FBtr0300430  NM_001169167   3835   995 br-RL FBtr0303562  NM_001201596   9620   514 br-RM FBtr0303563  NM_001201597   4213   724 br-RN FBtr0303564  NM_001201598   4546   724 br-RO FBtr0308319     4368   663
Additional Transcript Data & Comments
Reported size (kB)	11.5, 9.5, 7.4, 6.5, 4.5, 2 (northern blot) (Huang and Orr, 1992) 5.2 (northern blot) (Martinez-Laborda et al., 1992)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein br-PA   FBpp0070255   74.1   702   7.70     NP_726750   AAF45651 br-PB   FBpp0070253   92.3   880   7.98     NP_726752   AAF45647 br-PC   FBpp0070251   92.3   880   7.98     NP_726753   AAN09053 br-PE   FBpp0070252   77.1   724   7.87     NP_524759   AAF45648 br-PG   FBpp0070256   70.5   663   7.49     NP_726754   AAN09054 br-PH   FBpp0289656   92.3   880   7.98     NP_001138144   ACL82877 br-PI   FBpp0289657   92.3   880   7.98     NP_726751   AAF45650 br-PJ   FBpp0289658   55.5   514   7.12     NP_726749   AAN09052 br-PK   FBpp0289659   103.7   995   7.65     NP_001162638   ACZ95175 br-PL   FBpp0292595   55.5   514   7.12     NP_001188525   ADV37609 br-PM   FBpp0292596   77.1   724   7.87     NP_001188526   ADV37610 br-PN   FBpp0292597   77.1   724   7.87     NP_001188527   ADV37611 br-PO   FBpp0300638   70.5   663   7.49
Additional Polypeptide Data & Comments
Reported size (kDa)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • BTB/POZ-like (IPR000210) Zinc finger, C2H2 (IPR007087) BTB/POZ (IPR013069) BTB/POZ fold (IPR011333) Zinc finger, C2H2-like (IPR015880)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AE014298 AAF45647   AE014298 AAF45648   AE014298 AAF45650   AE014298 AAF45651   AE014298 AAN09052   AE014298 AAN09053   AE014298 AAN09054   AE014298 ACL82877   AE014298 ACZ95175   AE014298 ADV37609   AE014298 ADV37610   AE014298 ADV37611   AF345689   AF345690   AF345691   AF345692   AF345693   AF345694   AF345695   AF345696   AF345697   AF345698   AF345699   AF345700   AF345701   AI258226   AI260963   AI294595   AI296467   AI296538   AI297117   AI297782   AJ300017   AJ300018   AJ300019   AJ300020   AJ300021   AL009146 CAA15626   AL009146 CAA15627   AL009146 CAA15628   AL009146 CAA15629   AL021067   AW944129   AW944154   AY069756 AAL39901   BI573675   BI573676   BI616288   BI618664   BT125021   BZ748779   CA804100   CA806715   CA807496   CK657095   CK658663   CK659211   CK659720   CK660452   CK660638   CK661162   CO181559   CO182040   CO185750   CO186136   CO187594   CO188308   CO188403   CO192524   CO265347   CO267388   CO268177   CO269462   CO270073   CO271107   CO277445   CO277626   CO280396   CO280675   CO284573   CO284672   CO289558   CO289731   CO290682   CO293474   CO294721   CO296970   CO300385   CO305099   CO306404   CO309813   CO319118   CO319747   CO319772   CO320860   CO322061   CO323179   CO324246   CO325290   CO325797   CO327022   CO327220   CO328124   CO328652   CO329925   CO330589   CO331167   CO331212   CO332751   CO332864   CO333951   CO340386   CO341898   CO345288   CZ472859   CZ473454   CZ475289   CZ475290   CZ475657   CZ476127   DQ327739   EC059407   EC060279   EC060951   EC061426   EC061901   EC061921   EC063226   EC063834   EC065882   EC067059   EC071204   EC074231   EC074441   EC075574   EC076265   EC078537   EC079648   EC079977   EC081716   EC081783   EC084615   EC087004   EC090352   EC194392   EC197821   EC201851   EC206404   EC206548   EC208374   EC208463   EC211292   EC212134   EC214837   EC216137   EC216250   EC216630   EC216790   EC220107   EC221315   EC229039   EC232271   EC233335   EC233819   EC239018   EC240976   EC246830   EC250148   EC254009   EC266595   GH950306   GH950527   GH970945   GH973645   GH975588   GH983397   S55610   S55612   U51585 AAB09760   X54663 CAA38474   X54664 CAA38475   X54665 CAA38476   X54666 CAA38477   X62380   X63618   X97997   X97998   Z32013   Z32190   Z32387   Z32433
UniProtKB/Swiss-Prot	Q01295 Q24206
UniProtKB/TrEMBL	Q8IRX4 Q9W572 E1JJB2
Mapped Features
Mapped Features have been reorganized, please see this article for details. Additional mapped features and mutations can be found on GBrowse or related reports. Type Symbol & Location Additional Notes References
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
in situ Stage Tissue/Position (including subcellular localization) Reference oogenesis stage S5 -- S14 & adult stage | female   egg chamber   (Terashima and Bownes, 2004) northern blot Stage Tissue/Position (including subcellular localization) Reference late third instar larval stage   embryonic/larval central nervous system   (Restifo and Hauglum, 1998) embryonic/larval salivary gland   (Restifo and Hauglum, 1998) third instar - uncleared gut stage -- pupal stage P5(ii)   embryonic/larval central nervous system   (Restifo and Hauglum, 1998) adult stage   adult head   (Restifo and Hauglum, 1998) adult thorax & adult abdomen   (Restifo and Hauglum, 1998) Comment:described as "adult body" oogenesis & adult stage | female   ovary   (Huang and Orr, 1992, Martinez-Laborda et al., 1992) radioisotope in situ Stage Tissue/Position (including subcellular localization) Reference day 1 of adulthood -- day 2 of adulthood   adult brain   • neuronal cell body (Restifo and Hauglum, 1998)
Additional Descriptive Data
	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. (Huang and Orr, 1992) 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. (Woodard et al., 1994) 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. (Huang and Orr, 1992) 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. (Restifo and Hauglum, 1998) 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. (Huang and Orr, 1992) 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. (Terashima and Bownes, 2004)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
immunolocalization Stage Tissue/Position (including subcellular localization) Reference third instar larval stage   embryonic/larval salivary gland   • polytene chromosome (Gonzy et al., 2002) wandering third instar larval stage -- pupal stage P5(ii)   embryonic/larval central nervous system   • neuronal cell body (Restifo and Hauglum, 1998) embryonic/larval optic lobe   • neuronal cell body (Restifo and Hauglum, 1998) ventral nerve cord   • neuronal cell body (Restifo and Hauglum, 1998) prepupal stage P1 -- P3   salivary gland   • polytene chromosome (Gonzy et al., 2002) Comment:reference states 0-6 hr APF western blot Stage Tissue/Position (including subcellular localization) Reference late third instar larval stage   embryonic/larval central nervous system   (Restifo and Hauglum, 1998)
Additional Descriptive Data
	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. (Restifo and Hauglum, 1998, Gonzy et al., 2002)
Marker for
Subcellular Localization
CV Term
Notes
High-Throughput Expression Data
or
See Gelbart and Emmert, 2010.10.13 for analysis details and data files for all genes. modENCODE Temporal Expression Data for FBgn0000210    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0000210 Very low expression bin max Moderate expression bin max High expression bin max Extremely high expression bin max Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of high expression to a trough of extremely low expression. Peak expression observed during late larval stages, during early pupal stages. [download data (TSV)] Guide to modENCODE expression level colors   No expression (0 - 0)   Extremely low expression (1 - 10)   Very low expression (11 - 100)   Low expression (101 - 400)   Moderate expression (401 - 1400)   Moderately high expression (1401 - 4000)   High expression (4001 - 10000)   Very high expression (10001 - 100000)   Extremely high expression (100001 - 2000000) Linear, scaled to maximum FBgn0000210 expression level Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    24 embryo 04-06hr    13 embryo 06-08hr    36 embryo 08-10hr    55 embryo 10-12hr    100 embryo 12-14hr    172 embryo 14-16hr    381 embryo 16-18hr    343 embryo 18-20hr    358 embryo 20-22hr    240 embryo 22-24hr    201 larva L1    273 larva L2    84 larva L3 12hr old    210 larva L3 puffstage 1-2    1089 larva L3 puffstage 3-6    7135 larva L3 puffstage 7-9    9825 white prepupae new    6233 white prepupae 12hr    3165 white prepupae 24hr    919 pupae 2d postWPP    501 pupae 3d postWPP    120 pupae 4d postWPP    298 adult male 01day    82 adult male 05day    129 adult male 30day    128 adult female 01day    231 adult female 05day    219 adult female 30day    213 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    24 embryo 04-06hr    13 embryo 06-08hr    36 embryo 08-10hr    55 embryo 10-12hr    100 embryo 12-14hr  (172) embryo 14-16hr  (381) embryo 16-18hr  (343) embryo 18-20hr  (358) embryo 20-22hr  (240) embryo 22-24hr  (201) larva L1  (273) larva L2    84 larva L3 12hr old  (210) larva L3 puffstage 1-2  (1089) larva L3 puffstage 3-6  (7135) larva L3 puffstage 7-9  (9825) white prepupae new  (6233) white prepupae 12hr  (3165) white prepupae 24hr  (919) pupae 2d postWPP  (501) pupae 3d postWPP  (120) pupae 4d postWPP  (298) adult male 01day    82 adult male 05day  (129) adult male 30day  (128) adult female 01day  (231) adult female 05day  (219) adult female 30day  (213) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    24 embryo 04-06hr    13 embryo 06-08hr    36 embryo 08-10hr    55 embryo 10-12hr    100 embryo 12-14hr    172 embryo 14-16hr    381 embryo 16-18hr    343 embryo 18-20hr    358 embryo 20-22hr    240 embryo 22-24hr    201 larva L1    273 larva L2    84 larva L3 12hr old    210 larva L3 puffstage 1-2    1089 larva L3 puffstage 3-6  (7135) larva L3 puffstage 7-9  (9825) white prepupae new  (6233) white prepupae 12hr  (3165) white prepupae 24hr    919 pupae 2d postWPP    501 pupae 3d postWPP    120 pupae 4d postWPP    298 adult male 01day    82 adult male 05day    129 adult male 30day    128 adult female 01day    231 adult female 05day    219 adult female 30day    213 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    24 embryo 04-06hr    13 embryo 06-08hr    36 embryo 08-10hr    55 embryo 10-12hr    100 embryo 12-14hr    172 embryo 14-16hr    381 embryo 16-18hr    343 embryo 18-20hr    358 embryo 20-22hr    240 embryo 22-24hr    201 larva L1    273 larva L2    84 larva L3 12hr old    210 larva L3 puffstage 1-2    1089 larva L3 puffstage 3-6    7135 larva L3 puffstage 7-9    9825 white prepupae new    6233 white prepupae 12hr    3165 white prepupae 24hr    919 pupae 2d postWPP    501 pupae 3d postWPP    120 pupae 4d postWPP    298 adult male 01day    82 adult male 05day    129 adult male 30day    128 adult female 01day    231 adult female 05day    219 adult female 30day    213 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    24 embryo 04-06hr    13 embryo 06-08hr    36 embryo 08-10hr    55 embryo 10-12hr    100 embryo 12-14hr    172 embryo 14-16hr    381 embryo 16-18hr    343 embryo 18-20hr    358 embryo 20-22hr    240 embryo 22-24hr    201 larva L1    273 larva L2    84 larva L3 12hr old    210 larva L3 puffstage 1-2    1089 larva L3 puffstage 3-6    7135 larva L3 puffstage 7-9    9825 white prepupae new    6233 white prepupae 12hr    3165 white prepupae 24hr    919 pupae 2d postWPP    501 pupae 3d postWPP    120 pupae 4d postWPP    298 adult male 01day    82 adult male 05day    129 adult male 30day    128 adult female 01day    231 adult female 05day    219 adult female 30day    213 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0000210 expression level Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    24 embryo 04-06hr    13 embryo 06-08hr    36 embryo 08-10hr    55 embryo 10-12hr    100 embryo 12-14hr    172 embryo 14-16hr    381 embryo 16-18hr    343 embryo 18-20hr    358 embryo 20-22hr    240 embryo 22-24hr    201 larva L1    273 larva L2    84 larva L3 12hr old    210 larva L3 puffstage 1-2    1089 larva L3 puffstage 3-6    7135 larva L3 puffstage 7-9    9825 white prepupae new    6233 white prepupae 12hr    3165 white prepupae 24hr    919 pupae 2d postWPP    501 pupae 3d postWPP    120 pupae 4d postWPP    298 adult male 01day    82 adult male 05day    129 adult male 30day    128 adult female 01day    231 adult female 05day    219 adult female 30day    213 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    24 embryo 04-06hr    13 embryo 06-08hr    36 embryo 08-10hr    55 embryo 10-12hr    100 embryo 12-14hr  (172) embryo 14-16hr  (381) embryo 16-18hr  (343) embryo 18-20hr  (358) embryo 20-22hr  (240) embryo 22-24hr  (201) larva L1  (273) larva L2    84 larva L3 12hr old  (210) larva L3 puffstage 1-2  (1089) larva L3 puffstage 3-6  (7135) larva L3 puffstage 7-9  (9825) white prepupae new  (6233) white prepupae 12hr  (3165) white prepupae 24hr  (919) pupae 2d postWPP  (501) pupae 3d postWPP  120 pupae 4d postWPP  (298) adult male 01day    82 adult male 05day  129 adult male 30day  128 adult female 01day  (231) adult female 05day  (219) adult female 30day  (213) Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    24 embryo 04-06hr    13 embryo 06-08hr    36 embryo 08-10hr    55 embryo 10-12hr    100 embryo 12-14hr    172 embryo 14-16hr    381 embryo 16-18hr    343 embryo 18-20hr    358 embryo 20-22hr    240 embryo 22-24hr    201 larva L1    273 larva L2    84 larva L3 12hr old    210 larva L3 puffstage 1-2    1089 larva L3 puffstage 3-6  (7135) larva L3 puffstage 7-9  (9825) white prepupae new  (6233) white prepupae 12hr  (3165) white prepupae 24hr    919 pupae 2d postWPP    501 pupae 3d postWPP    120 pupae 4d postWPP    298 adult male 01day    82 adult male 05day    129 adult male 30day    128 adult female 01day    231 adult female 05day    219 adult female 30day    213 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    24 embryo 04-06hr    13 embryo 06-08hr    36 embryo 08-10hr    55 embryo 10-12hr    100 embryo 12-14hr    172 embryo 14-16hr    381 embryo 16-18hr    343 embryo 18-20hr    358 embryo 20-22hr    240 embryo 22-24hr    201 larva L1    273 larva L2    84 larva L3 12hr old    210 larva L3 puffstage 1-2    1089 larva L3 puffstage 3-6    7135 larva L3 puffstage 7-9    9825 white prepupae new    6233 white prepupae 12hr    3165 white prepupae 24hr    919 pupae 2d postWPP    501 pupae 3d postWPP    120 pupae 4d postWPP    298 adult male 01day    82 adult male 05day    129 adult male 30day    128 adult female 01day    231 adult female 05day    219 adult female 30day    213 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    24 embryo 04-06hr    13 embryo 06-08hr    36 embryo 08-10hr    55 embryo 10-12hr    100 embryo 12-14hr    172 embryo 14-16hr    381 embryo 16-18hr    343 embryo 18-20hr    358 embryo 20-22hr    240 embryo 22-24hr    201 larva L1    273 larva L2    84 larva L3 12hr old    210 larva L3 puffstage 1-2    1089 larva L3 puffstage 3-6    7135 larva L3 puffstage 7-9    9825 white prepupae new    6233 white prepupae 12hr    3165 white prepupae 24hr    919 pupae 2d postWPP    501 pupae 3d postWPP    120 pupae 4d postWPP    298 adult male 01day    82 adult male 05day    129 adult male 30day    128 adult female 01day    231 adult female 05day    219 adult female 30day    213 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Developmental Stage   Expression Level embryo 00-02hr     embryo 02-04hr     embryo 04-06hr     embryo 06-08hr     embryo 08-10hr     embryo 10-12hr     embryo 12-14hr     embryo 14-16hr     embryo 16-18hr     embryo 18-20hr     embryo 20-22hr     embryo 22-24hr     larva L1     larva L2     larva L3 12hr old     larva L3 puffstage 1-2     larva L3 puffstage 3-6     larva L3 puffstage 7-9     white prepupae new     white prepupae 12hr     white prepupae 24hr     pupae 2d postWPP     pupae 3d postWPP     pupae 4d postWPP     adult male 01day     adult male 05day     adult male 30day     adult female 01day     adult female 05day     adult female 30day     FlyAtlas Anatomical Expression Data for FBgn0000210    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0000210 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Two or more Affy2 ProbeSets identify exons of this gene. This is a summary of the tissue expression peaks exhibited in at least one of these ProbeSets. Expression at high levels in the following post-embryonic organs or tissues: larval salivary gland. Expression at moderate levels in the following post-embryonic organs or tissues: larval/adult central nervous system, larval hindgut, larval Malpighian tubules, larval fat body, larval trachea, larval carcass. [download data (TSV)] Guide to FlyAtlas expression level colors   No expression (0 - 9.999)   Low expression (10 - 99.999)   Moderate expression (100 - 499.999)   High level expression (500 - 999.999)   Very high expression (1000 - 25000) Heatmap Tissue   Expression Level Larval Central Nervous System no informative data Larval Midgut no informative data Larval Hindgut no informative data Larval Malpighian Tubules no informative data Larval Fat Body no informative data Larval Salivary Gland no informative data Larval Trachea no informative data Larval Carcass no informative data Adult Head     Adult Eye     Adult Brain no informative data Adult Thoracic-Abdominal Ganglion no informative data Adult Crop     Adult Midgut     Adult Hindgut     Adult Malpighian Tubules     Adult Fat Body no informative data Adult Salivary Gland     Adult Heart     Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca     Adult Ovary no informative data Adult Testis     Adult Male Accessory Gland     Adult Carcass     modENCODE Temporal Expression Data (Graveley et al., 2011) FlyAtlas Anatomical Expression Data (Chintapalli et al., 2007)
Expression Clusters
	A cluster of genes with similar mRNA expression dynamics across development. (The modENCODE Consortium, 2010) mE2_34_mRNA_expression_cluster_25
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0000210 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG11491-RC;CG11491-RA;CG11491-RF;CG11491-RD;CG11491-RE FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for br Stages(s) 13-16
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal brnpr-6 brnpr-7 lethal, with Scer\GAL4Feb211 brZ3.Scer\UAS lethal, with Scer\GAL4P0206 brZ3.Scer\UAS lethal (with br6) br7 lethal (with br7) br6 lethal (with brnpr-5) brrbp-2 lethal (with brnpr-6) brrbp-2 brrbp-5 lethal (with brrbp-2) brnpr-5 brnpr-6 lethal (with brrbp-5) brnpr-6 lethal (with Df(1)br25) brnpr-5 brnpr-6 lethal | adult stage brrbp-1 lethal | embryonic stage, with Scer\GAL4elav-C155 brZ3.Scer\UAS lethal | female (with br6) brnpr-6 lethal | female (with brnpr-6) br6 lethal | first instar larval stage, with Scer\GAL4elav-C155 brZ3.Scer\UAS lethal | first instar larval stage | heat sensitive brBRcore.NS.Z3.hs lethal | heat sensitive brBRcore.NS.Z3.hs brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs brBRcore.Z4.hs brZ2.hs lethal | larval stage brG0018 brG0042 brnpr-1 brnpr-3 brnpr-4 brnpr-6 lethal | larval stage, with Scer\GAL4dpp.blk1 brZ2.Scer\UAS lethal | larval stage (with br5) brnpr-6 lethal | larval stage (with brnpr-6) br5 lethal | larval stage (with Df(1)P154) brnpr-6 lethal | larval stage | heat sensitive brBRcore.Z4.hs brZ2.hs lethal | larval stage | recessive br18 br23RL br30OM br38JL br80BL brnpr-1 brnpr-3 brnpr-5 brnpr-6 brnpr-7 brnpr-M8 lethal | maternal effect br20 lethal | partially | pharate adult stage (with brnpr-3) brrbp-2 lethal | partially | pharate adult stage (with brrbp-2) brnpr-3 lethal | partially | pupal stage (with brnpr-3) brrbp-2 lethal | partially | pupal stage (with brrbp-2) brnpr-3 lethal | partially | recessive br6 brrbp-2 lethal | pharate adult stage br2Bd brrbp-1 lethal | pharate adult stage, with Scer\GAL4Feb204 brZ3.Scer\UAS lethal | pharate adult stage | recessive brrbp-1 lethal | pharate adult stage P8 | heat sensitive brBRcore.NS.Z3.hs lethal | prepupal stage br2Bc-1 br5 br28 brG0018 brG0042 brG0401 lethal | prepupal stage | recessive br2Bab-8 br2Bab-9 br2Bab-10 br2Bab-M3 br2Bab-M4 br2Bab-M7 br2Bc-1 br5 br7 br8 br25.4 br28 br29 brBC1 lethal | pupal stage br2Bc-1 br6 brG0318 brnpr-1 lethal | pupal stage, with Scer\GAL4pnr-MD237 brGD4279 lethal | pupal stage (with br2Bc-2) brnpr-6 lethal | pupal stage (with brnpr-6) br2Bc-2 lethal | pupal stage | recessive br2Bab-1 br2Bab-2 br2Bab-4 br2Bc-2 br6 brBC3 brnpr-1 brnpr-2 brrbp-1 brrbp-3 brrbp-4 brrbp-5 lethal | pupal stage P7, with Scer\GAL4Feb204 brZ3.Scer\UAS lethal | recessive br2Bab-1 br2Bab-2 br2Bab-3 br2Bab-8 br2Bab-M2 br2Bc-1 br2Bc-2 br5 br7 br8 br10 br11 br12 br13 br14 br15 br16 br17 br18 br20 br21 br22 br23 br24 br28 br29 brD brESHS5 brG0018 brG0042 brG0318 brG0401 brl-a brl-b brnpr-3 brPG12 brPG83 brPL86 brrbp-1 brrbp-4 brrbp-5 lethal | recessive | cold sensitive br23 br24 lethal | recessive | heat sensitive br2Bd br5 lethal | second instar larval stage | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs lethal | second instar larval stage | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs lethal | third instar larval stage, with Scer\GAL4P0206 brZ4.Scer\UAS semi-lethal | pupal stage brrbp-1 semi-lethal | pupal stage, with Scer\GAL4pnr-MD237 brGD7318 viable br1 br2Bab-7 br2Bd br3 br4 br9 br24h br27 br33a br37l brrbp-1 brrbp-2 brsh bruq brw viable (with Df(1)br25) brrbp-2 brrbp-3 brrbp-4 brrbp-5 viable | good bruq viable | reduced (with Df(1)br25) br6 br7 brnpr-6 Sterility Allele female fertile br1 female sterile (with brnpr-6) brrbp-1 brrbp-2 female sterile (with brrbp-1) brnpr-6 female sterile (with brrbp-2) brnpr-6 female sterile | recessive brnpr-fs brrbp-4 fertile brrbp-2 bruq fertile | good bruq male fertile br1 Other Phenotypes Allele body color defective | heat sensitive brBRcore.NS.Z3.hs brBRcore.TNT.Q1.Z1.hs brBRcore.Z4.hs body color defective | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs body color defective | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs chemical resistant brhardy decreased cell death brdsRNA.cKa brrbp-5 developmental rate defective | second instar larval stage br1 eye color defective br2Bab-M7 flightless | partially brrbp-2 immune response defective brnpr-3 locomotor behavior defective | first instar larval stage, with Scer\GAL4elav-C155 brZ3.Scer\UAS locomotor behavior defective | second instar larval stage, with Scer\GAL4elav-C155 brZ3.Scer\UAS long lived | male brhardy Minute brnpr-3 neuroanatomy defective | adult stage, with Scer\GAL4P2.4.Pdf brZ3.Scer\UAS visible, with Scer\GAL4pnr-MD237 brGD7318 visible (with br2Bc-1) br25.4 brBC3 visible (with br25.4) br2Bc-1 visible (with brBC3) br2Bc-1 visible (with T(1;2)dorvar7) brnpr-6 visible | heat sensitive br1 br3 brBRcore.TNT.Q1.Z1.hs visible | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs visible | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs visible | recessive brR1 bruq visible | recessive | cold sensitive br23 br24 visible | recessive | somatic clone brnpr-3 wild-type brBRcore.NS.Z3.hs brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs Phenotype manifest in Allele abdomen br2Bd brrbp-2 abdominal dorsal trichome | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs abdominal dorsal trichome | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs abdominal sternite bristle brrbp-5 adult abdominal sensillum | heat sensitive brBRcore.TNT.Q1.Z1.hs adult abdominal sensillum | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs adult abdominal sensillum | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs adult brain br1 br27 br2Bc-1 br2Bc-2 br5 br6 brrbp-1 brrbp-4 adult cuticle | heat sensitive brBRcore.TNT.Q1.Z1.hs adult cuticle | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs adult cuticle | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs adult epidermis br5 brrbp-5 adult head br5 adult head (with br2Bc-1) br25.4 adult head (with br25.4) br2Bc-1 chaeta | heat sensitive brBRcore.NS.Z3.hs brBRcore.Z4.hs chemosensory ventral triple row bristle | somatic clone brnpr-3 chorion brnpr-fs brrbp-1 chorion | cold sensitive brBRcore.NS.Z3.hs coxal tergal remotor muscle brrbp-1 brrbp-2 brrbp-4 brrbp-5 dorsal appendage brnpr-fs brrbp-1 brrbp-2 dorsal appendage (with br1) brA47 brnpr-6 brrbp-1 dorsal appendage (with brA47) br1 dorsal appendage (with brnpr-6) br1 brrbp-2 dorsal appendage (with brrbp-1) br1 dorsal appendage (with brrbp-2) brnpr-6 dorsal appendage | ectopic | cold sensitive brBRcore.NS.Z3.hs brBRcore.TNT.Q1.Z1.hs brBRcore.Z4.hs brZ2.hs dorsal appendage | germline clone br20 dorsal appendage | somatic clone brCJ89 dorsal medial muscle brrbp-1 brrbp-4 brrbp-5 egg | germline clone br20 egg operculum | somatic clone brCJ89 embryonic/larval central nervous system brnpr-3 embryonic/larval esophagus | prepupal stage br2Bc-2 embryonic/larval midgut | prepupal stage br2Bc-2 embryonic/larval proventriculus | embryonic stage br1 br27 br2Bc-1 br2Bc-2 br5 br6 brrbp-1 brrbp-4 embryonic/larval proventriculus | prepupal stage br2Bc-2 embryonic/larval salivary gland brrbp-5 embryonic/larval salivary gland | embryonic stage br1 br27 br2Bab-1 br2Bc-1 br2Bc-2 br5 br6 brnpr-3 brnpr-7 brrbp-1 brrbp-4 brrbp-5 embryonic/larval somatic muscle brrbp-5 endochorion brnpr-fs eye brrbp-2 brrbp-5 bruq eye | somatic clone br5 brnpr-3 follicle cell | cold sensitive brBRcore.TNT.Q1.Z1.hs head bristle | heat sensitive brBRcore.TNT.Q1.Z1.hs head bristle | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs head bristle | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs imaginal disc br29 br8 brnpr-3 imaginal disc | larval stage brnpr-3 brnpr-4 indirect flight muscle br1 br27 br2Bab-1 br2Bc-1 br2Bc-2 br5 br6 brnpr-3 brnpr-7 brrbp-1 brrbp-4 lateral oblique dorsal muscle brrbp-1 brrbp-2 brrbp-4 brrbp-5 leg br1 br23 br24 br2Bab-1 br5 brsh leg | P-stage brrbp-5 LNv neuron, with Scer\GAL4P2.4.Pdf brZ3.Scer\UAS lobula br2Bc-2 lobula plate br2Bc-2 br6 macrochaeta br23 br24 bruq macrochaeta | heat sensitive brBRcore.TNT.Q1.Z1.hs macrochaeta | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs macrochaeta | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs male genitalia br2Bc-2 brrbp-1 male genitalia, with br5 brZ2.hs male genitalia, with brZ2.hs br5 male genitalia (with Dp(1;Y)Sz280) brrbp-1 Malpighian tubule br2Bc-1 br2Bc-2 maxillary palp bristle br2Bab-1 br2Bab-2 br2Bd brR1 brnpr-7 brrbp-1 brrbp-2 brrbp-5 maxillary palp bristle (with br6) brrbp-3 brrbp-4 maxillary palp bristle (with brnpr-5) brrbp-2 maxillary palp bristle (with brnpr-6) brrbp-2 brrbp-5 maxillary palp bristle (with brrbp-2) brnpr-5 brnpr-6 maxillary palp bristle (with brrbp-3) br6 maxillary palp bristle (with brrbp-4) br6 maxillary palp bristle (with brrbp-5) brnpr-6 maxillary palp bristle (with Df(1)br25) brrbp-2 brrbp-3 brrbp-4 brrbp-5 maxillary palpus br1 br6 br7 maxillary palpus & macrochaeta br1 br2Bab-2 brrbp-2 medulla br2Bc-2 br6 mesothoracic bristle | heat sensitive brBRcore.TNT.Q1.Z1.hs mesothoracic bristle | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs mesothoracic bristle | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs mesothoracic extracoxal depressor muscle 66 brrbp-1 brrbp-4 brrbp-5 mesothoracic tergum, with Scer\GAL4pnr-MD237 brGD7318 metathoracic bristle | heat sensitive brBRcore.TNT.Q1.Z1.hs metathoracic bristle | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs metathoracic bristle | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs metathoracic leg (with br2Bc-1) br25.4 brBC3 metathoracic leg (with br25.4) br2Bc-1 metathoracic leg (with brBC3) br2Bc-1 metathoracic tarsal segment (with br2Bab-1) br2Bc-1 metathoracic tarsal segment (with br2Bc-1) br2Bab-1 microchaeta & adult thorax bruq microchaeta & thorax bruq microchaeta | heat sensitive brBRcore.TNT.Q1.Z1.hs microchaeta | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs microchaeta | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs micropyle | cold sensitive brBRcore.NS.Z3.hs morphogenetic furrow br5 brnpr-1 brnpr-3 morphogenetic furrow | somatic clone br5 muscle tendon junction & coxal tergal remotor muscle brrbp-1 muscle tendon junction & lateral oblique dorsal muscle brrbp-1 muscle tendon junction & tergosternal muscle brrbp-1 ommatidium br5 brnpr-3 ommatidium | somatic clone br5 brnpr-3 optic lobe br1 br27 br2Bc-1 br2Bc-2 br5 br6 brrbp-1 brrbp-4 ovary br1 brrbp-1 brrbp-2 ovary (with brnpr-6) brrbp-2 ovary (with brrbp-2) brnpr-6 ovary & nucleus | conditional cs brBRcore.TNT.Q1.Z1.hs ovary | cold sensitive brBRcore.TNT.Q1.Z1.hs photoreceptor cell brnpr-1 photoreceptor cell | somatic clone br5 brnpr-3 photoreceptor cell | somatic clone | supernumerary br5 brnpr-3 photoreceptor cell R8 | somatic clone | supernumerary br5 brnpr-3 pigment cell br2Bab-M7 polytene chromosome puff br2Bbc-1 prothoracic gland, with Scer\GAL4P0206 brZ1.Scer\UAS brZ2.Scer\UAS brZ3.Scer\UAS pupa brG0318 puparium br5 ring gland brnpr-4 rostral membrane br1 brnpr-7 salivary gland brrbp-5 scutellar bristle brrbp-2 sensory neuron & dorsal mesothoracic disc, with Scer\GAL4bbg-C96 brZ1.Scer\UAS socket | heat sensitive brBRcore.Z4.hs tendon cell brrbp-1 tergite | heat sensitive, with brBRcore.Q1.Z1.hs brBRcore.TNT.Q1.Z1.hs tergite | heat sensitive, with brBRcore.TNT.Q1.Z1.hs brBRcore.Q1.Z1.hs tergosternal muscle brrbp-1 brrbp-2 brrbp-4 brrbp-5 thorax brrbp-1 trichogen cell, with Scer\GAL4pnr-MD237 brGD7318 ventral thoracic disc br5 wing br1 br23 br24 br2Bab-1 br2Bab-2 br2Bd br3 br6 brD brR1 brnpr-6 brrbp-2 brsh bruq wing (with br2Bc-1) br25.4 wing (with br6) br7 brrbp-3 brrbp-4 wing (with br7) br6 wing (with br25.4) br2Bc-1 wing (with brrbp-3) br6 wing (with brrbp-4) br6 wing (with Df(1)br25) br6 br7 brnpr-6 wing (with T(1;2)dorvar7) brnpr-6 wing | heat sensitive brBRcore.TNT.Q1.Z1.hs wing margin & sensory neuron, with Scer\GAL4bbg-C96 brZ1.Scer\UAS wing margin & sensory neuron | somatic clone brnpr-3 yellow body br2Bc-1 br2Bc-2
Classical Alleles ( 118 )
For All Classical Alleles Show
Allele of br Class Mutagen Stocks Known lesion br1 hypomorphic allele - genetic evidence spontaneous 9 Yes brnpr-3 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate 3 -- br3 hypomorphic allele - genetic evidence spontaneous 2 Yes brG0284a P-element activity 2 -- brG0318 P-element activity 2 -- brG0401 P-element activity 2 -- br28 amorphic allele - genetic evidence, loss of function allele PM hybrid dysgenesis P-element activity 1 Yes br2Bab-5 amorphic allele - genetic evidence diepoxybutane 1 Yes br2Bc-1 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate 1 -- br2Bc-2 amorphic allele - genetic evidence, hypomorphic allele - genetic evidence ethyl methanesulfonate 1 -- br5 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate 1 -- brnpr-5 amorphic allele - genetic evidence ethyl methanesulfonate 1 Yes brnpr-6 amorphic allele - genetic evidence ethyl methanesulfonate 1 -- br2Bc-4 ethyl methanesulfonate 1 -- brc00809 1 -- brd03598 P-element activity 1 -- brd06704 P-element activity 1 -- brEP1515 P-element activity 1 -- brEY00662 P-element activity 1 -- brG0018 P-element activity 1 -- brG0042 P-element activity 1 -- brGS1116 1 Yes brKG04048 1 -- brKG05923 1 -- brNP4070 1 -- brNP6018 1 -- brnpr-1 ethyl methanesulfonate 1 -- brrbp-1 hypomorphic allele - genetic evidence ethyl methanesulfonate 1 -- brrbp-4 hypomorphic allele - genetic evidence ethyl methanesulfonate 1 -- brrbp-5 loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate 1 Yes br2Bab-1 amorphic allele - genetic evidence, hypomorphic allele - genetic evidence ethyl methanesulfonate 0 -- br8 amorphic allele - genetic evidence ethyl methanesulfonate 0 -- brnpr-7 loss of function allele, amorphic allele - genetic evidence diepoxybutane 0 -- brnpr-x1002 amorphic allele - genetic evidence X ray 0 Yes br10 X ray 0 -- br11 X ray 0 -- br12 X ray 0 -- br13 X ray 0 -- br14 X ray 0 -- br15 X ray 0 -- br16 X ray 0 -- br17 X ray 0 -- br18 ethyl methanesulfonate 0 -- br1U 0 Yes br20 ethyl methanesulfonate 0 -- br21 ethyl methanesulfonate 0 -- br22 ethyl methanesulfonate X ray 0 -- br23 spontaneous 0 -- br23RL Delta2-3 0 Yes br24 spontaneous 0 -- br24h spontaneous 0 -- br25.4 P-element activity 0 Yes br25 loss of function allele X ray 0 Yes br26 loss of function allele, hypomorphic allele - genetic evidence diepoxybutane 0 -- br27 hypomorphic allele - genetic evidence diepoxybutane 0 -- br29 X ray 0 Yes br2Bab-10 gene conversion Delta2-3 0 Yes br2Bab-2 ethyl methanesulfonate 0 -- br2Bab-3 ethyl methanesulfonate 0 -- br2Bab-4 0 -- br2Bab-6 0 -- br2Bab-7 P-element activity 0 Yes br2Bab-8 gene conversion Delta2-3 0 Yes br2Bab-9 gene conversion Delta2-3 0 Yes br2Bab-M2 gene conversion Delta2-3 0 Yes br2Bab-M3 gene conversion Delta2-3 0 Yes br2Bab-M4 gene conversion Delta2-3 0 Yes br2Bab-M7 gene conversion Delta2-3 0 Yes br2Bbc-1 gene conversion Delta2-3 0 Yes br2Bc-3 ethyl methanesulfonate 0 -- br2Bc-6 PM hybrid dysgenesis P-element activity 0 Yes br2Bd hypomorphic allele - genetic evidence ethyl methanesulfonate 0 -- br30OM P-element activity 0 Yes br33a spontaneous 0 -- br37l spontaneous 0 -- br38JL Delta2-3 0 Yes br45j natural population 0 -- br4 DNA 0 Yes br5-HA-2643 0 -- br6 loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes br7 ethyl methanesulfonate 0 -- br80BL Delta2-3 0 Yes br9 hypomorphic allele - genetic evidence gamma ray 0 -- brA47 P-element activity 0 -- brAub hypomorphic allele - genetic evidence 0 -- brBC1 Delta2-3 0 Yes brBC3 Delta2-3 0 Yes brCJ89 ethyl methanesulfonate 0 -- brd02514 P-element activity 0 -- brd07313 P-element activity 0 -- brd07726 0 -- brd08215 P-element activity 0 -- brd10457 0 -- brD spontaneous 0 -- brecs-lt76 0 -- brESHS5 MRh12 MR 0 Yes brGG01056 0 -- brGG01780 0 -- brhardy P-element activity 0 Yes brl-a spontaneous 0 -- brl-b spontaneous 0 -- brnpr-2 ethyl methanesulfonate 0 -- brnpr-4 ethyl methanesulfonate 0 -- brnpr-fs P-element activity PM hybrid dysgenesis 0 Yes brnpr-M8 gene conversion Delta2-3 0 Yes brPG12 Delta2-3 0 -- brPG83 Delta2-3 0 -- brPL86 Delta2-3 0 -- brR1 0 -- brrbp-2 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 -- brrbp-3 ethyl methanesulfonate 0 -- brrbp-lp13 0 -- brsh spontaneous 0 -- brunspecified 0 -- bruq2 X ray 0 -- bruq p-N,N-di-(2-chloroethyl)amino-D-phenylalanine 0 -- brw spontaneous 0 -- brZ4.hs 0 --
Alleles Carried on Transgenic Constructs ( 23 )
For All Alleles Carried on Transgenic Constructs Show
Allele of br Class Mutagen Stocks Known lesion brGD4279 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes brGD7318 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes brHMS00042 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes brJF02585 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes brKK111704 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes brBRcore.NS.Z3.hs in vitro construct - regulatory fusion 0 Yes brBRcore.Q1.Z1.hs in vitro construct - regulatory fusion 0 Yes brBRcore.TNT.Q1.Z1.hs in vitro construct - regulatory fusion 0 Yes brBRcore.Z2.hs in vitro construct - regulatory fusion 0 Yes brBRcore.Z4.hs in vitro construct - regulatory fusion 0 Yes brdsRNA.392-1060 in vitro construct - RNAi 0 Yes brdsRNA.cKa in vitro construct - RNAi 0 Yes brdsRNA.cSa in vitro construct - RNAi 0 Yes brGD4278 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes brZ1.IE in vitro construct - regulatory fusion 0 Yes brZ1.Scer\UAS in vitro construct - regulatory fusion 0 Yes brZ2.hs in vitro construct - regulatory fusion 0 Yes brZ2.IE in vitro construct - regulatory fusion 0 Yes brZ2.Scer\UAS in vitro construct - regulatory fusion 0 Yes brZ2.T:Zzzz\FLAG in vitro construct - coding region fusion 0 Yes brZ3.IE in vitro construct - regulatory fusion 0 Yes brZ3.Scer\UAS in vitro construct - regulatory fusion 0 Yes brZ4.Scer\UAS in vitro construct - regulatory fusion 0 Yes
Aneuploid Aberrations
Duplicated in	Dp(1;1)100 (Dobzhansky, 1932) Dp(1;1)105 (Dobzhansky, 1932) Dp(1;3)sta (Kramers et al., 1983, Kramers et al., 1991) Dp(1;f)100 (Morgan, 1938) Dp(1;f)102 (Dobzhansky, 1932) Dp(1;f)136 (Dobzhansky, 1932) Dp(1;f)137 (Dobzhansky, 1932) Dp(1;f)R (Eeken et al., 1985) Dp(1;Y)2E (Aizenzon and Belyaeva, 1982) Dp(1;Y)59k9-4 (Kramers et al., 1983, Kramers et al., 1991) Dp(1;Y)Sz280 (Belyaeva et al., 1987) Dp(1;Y)y221T (Belyaeva et al., 1980) Dp(1;Y)y240T (Belyaeva et al., 1980) Dp(1;Y)y253T (Belyaeva et al., 1980, Aizenzon and Belyaeva, 1982) Dp(1;Y)y260d19.3   Dp(1;Y)y267g19.1 (Liu and Restifo, 1998, Belyaeva et al., 1980, Belyaeva et al., 1981, Lefevre, 1981, Aizenzon and Belyaeva, 1982, Eeken et al., 1985, Schalet, 1986, Restifo and White, 1991) Dp(1;Y)y267g24.2 (Belyaeva et al., 1980, Aizenzon and Belyaeva, 1982, Protopopov et al., 1991)
Disrupted in	Df(1)A52.3 (Prudhommeau and Proust, 1990) Df(1)A94 (Craymer and Roy, 1980, Kramers et al., 1983, Eeken et al., 1985, Kramers et al., 1991) Df(1)br25 (Belyaeva et al., 1980, Belyaeva et al., 1982, Belyaeva et al., 1987, Mazina et al., 1991) Df(1)br-rv1 (Belyaeva et al., 1987) Df(1)BSC589 (Cook et al., 2008.9.3) Df(1)BSC719 (Christensen et al., 2008.12.28) Df(1)Exel8196 (Ryder, 2004.4.26) Df(1)H27.2 (Prudhommeau and Proust, 1990) Df(1)JTΔ1 (Mazina et al., 1991) Df(1)P154 (Belyaeva et al., 1987, Mazina et al., 1991, Shestopal, 1997.2.6) Df(1)RA19 (Belyaeva et al., 1980, Belyaeva et al., 1982, Aizenzon and Belyaeva, 1982) Df(1)S39 (Liu and Restifo, 1998, Craymer and Roy, 1980, Belyaeva et al., 1981, Belyaeva et al., 1982, Kramers et al., 1983, Belyaeva et al., 1987, Restifo and White, 1991, Kramers et al., 1991, Lehmann and Korge, 1996) Df(1)Sp(br-dor) (Schalet, 1986) Df(1)St469 (Belyaeva et al., 1987) Df(1)sta (Kramers et al., 1983, Kramers et al., 1991) Dp(1;Y)Sz280 (Wilson et al., 2006) In(1)br103 (Belyaeva et al., 1987, Mazina et al., 1991) In(1)Hw49c (Mazina et al., 1991) In(1)l(1)2Bab5 (Gonzy-Treboul et al., 1995) T(1;2)br4 (Protopopov et al., 1991, Mazina et al., 1991) T(1;3)JTΔ3 (Protopopov et al., 1991, Mazina et al., 1991) T(1;3)npr1 (O'Keefe et al., 1995) T(1;4)231b (Patterson, 1932) Ts(1Lt;YLt)G20 (Belyaeva et al., 1987) Ts(1Lt;YSt)B131 (Belyaeva et al., 1987)
Not duplicated in	Dp(1;1)112 (Dobzhansky, 1932) Dp(1;3)sta (Melnick et al., 1993) Dp(1;f)101 (Belyaeva et al., 1980, Aizenzon and Belyaeva, 1982, Melnick et al., 1993) Dp(1;f)106 (Dobzhansky, 1932) Dp(1;f)107 (Dobzhansky, 1932) Dp(1;f)118 (Dobzhansky, 1932) Dp(1;Y)dor21T (Belyaeva et al., 1982) Dp(1;Y)Sz280 (Liu and Restifo, 1998, Belyaeva et al., 1981, Belyaeva et al., 1982, Aizenzon and Belyaeva, 1982, Protopopov et al., 1991, Restifo and White, 1991, Lehmann and Korge, 1996) Dp(1;Y)y59b (Belyaeva et al., 1980, Aizenzon and Belyaeva, 1982) Dp(1;Y)y+mal+ (Eeken et al., 1985)
Not disrupted in	Df(1)247g (Patterson, 1932) Df(1)262 (Patterson, 1932) Df(1)267 (Patterson, 1932) Df(1)303 (Patterson, 1932) Df(1)308 (Patterson, 1932) Df(1)314 (Patterson, 1932) Df(1)403 (Kramers et al., 1991) Df(1)At127 (Aizenzon and Belyaeva, 1982) Df(1)cin-arth (Kramers et al., 1991) Df(1)HA18 (Schalet, 1986) Df(1)RA19 (Belyaeva et al., 1982) Df(1)S39 (Aizenzon and Belyaeva, 1982) Df(1)Sp(J1a) (Schalet, 1986) Df(1)Sp(J1b) (Schalet, 1986) Df(1)Sp(J1c) (Schalet, 1986) Df(1)St472 (Aizenzon and Belyaeva, 1982, Belyaeva et al., 1987) Df(1)St490 (Belyaeva et al., 1987, Mazina et al., 1991) Df(1)sta (Belyaeva et al., 1982, Aizenzon and Belyaeva, 1982, Eeken et al., 1985, Belyaeva et al., 1987, Mazina et al., 1991, Melnick et al., 1993) Df(1)svr (Kramers et al., 1991) Df(1)y74k24.1 (Eeken et al., 1985, Kramers et al., 1991)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD4278} NA P{GD4279} NA P{GD7318} NA P{KK111704} NA P{TRiP.HMS00042} NA P{TRiP.JF02585} NA P{UAS-br.Z1} NA P{UAS-br.Z2} NA P{UAS-br.Z4} NA heat-shock construct P{BRcore-Q1-Z1} NA P{BRcore-Z2} NA P{hs-BRC-Z1} NA P{hs-BRC-Z2} NA P{hs-BRC-Z3} NA P{hs-BRC-Z4} NA
Insertions
	Type of insertions Name Expression data insertion of mobile activating element P{EP}brEP1515 NA P{EPgy2}brEY00662 NA P{GSV1}brGS1116 NA P{Mae-UAS.6.11}B0731B0731 NA P{Mae-UAS.6.11}B2005B2005 NA P{Mae-UAS.6.11}brGG01056 NA P{Mae-UAS.6.11}brGG01780 NA P{Mae-UAS.6.11}GG01083GG01083 NA P{XP}brd02514 NA P{XP}brd03598 NA P{XP}brd06704 NA P{XP}brd07313 NA P{XP}brd07726 NA P{XP}brd08215 NA P{XP}brd10457 NA insertion of enhancer trap binary system P{GAL4}brA47 No P{GAL4}brhardy No P{GawB}brNP4070 No P{GawB}brNP6018 No P{GawB}brPG12 No P{GawB}brPG83 No miscellaneous insertions 412{}br1 NA 412{}br3b NA gypsy{}brnpr-fs NA P{3'lacW.2Bab-10}br2Bab-10 NA P{3'lacW.2Bbc-1}br30OM NA P{5'lacW}br38JL NA P{}br2Bab-7 NA P{}br2Bc-6 NA P{}br28 NA P{lacW.2Bbc-1}br2Bbc-1 NA P{lacW-Δ80BL}br80BL NA P{RS5}br5-HA-2643 NA P{SUPor-P}brKG04048 NA P{SUPor-P}brKG05923 NA PBac{PB}brc00809 NA roo{}br3a NA insertion of enhancer trap P{lacW}br2Bab-8 No P{lacW}br2Bab-9 No P{lacW}br2Bab-M2 No P{lacW}br2Bab-M3 No P{lacW}br2Bab-M4 No P{lacW}br23RL No P{lacW}brG0018 No P{lacW}brG0042 No P{lacW}brG0284a No P{lacW}brG0318 No P{lacW}brG0401 No P{lacW}brPL86 No
Gene Ontology: Function, Process & Cellular Component ( 25 unique terms )
Terms Based on Experimental Evidence ( 18 terms )
Molecular Function
CV term Evidence References
DNA binding inferred from direct assay (Chen et al., 2002) RNA polymerase II regulatory region sequence-specific DNA binding inferred from direct assay (Crossgrove et al., 1996)
Biological Process
CV term Evidence References
autophagy inferred from mutant phenotype (Lee and Baehrecke, 2001) cellular response to ecdysone inferred from mutant phenotype (Daish et al., 2003) compound eye photoreceptor fate commitment inferred from mutant phenotype (Brennan et al., 2001) eclosion inferred from expression pattern (Chen et al., 2002) larval central nervous system remodeling inferred from mutant phenotype (Spokony and Restifo, 2003) male genitalia morphogenesis inferred from mutant phenotype (Wilson et al., 2006) muscle fiber development inferred from genetic interaction with Mef2 (Lovato et al., 2005) inferred from genetic interaction with twi (Lovato et al., 2005) negative regulation of gene expression inferred from mutant phenotype (Ward et al., 2006) oogenesis inferred from mutant phenotype (Wilson et al., 2006) positive regulation of transcription from RNA polymerase II promoter inferred from mutant phenotype (Daish et al., 2003, Crossgrove et al., 1996) progression of morphogenetic furrow involved in compound eye morphogenesis inferred from mutant phenotype (Brennan et al., 1998, Brennan et al., 2001) regulation of development, heterochronic inferred from mutant phenotype (Parrish et al., 2009) response to ecdysone inferred from mutant phenotype (Sempere et al., 2002) response to symbiont inferred from mutant phenotype (Sorrentino et al., 2002) salivary gland histolysis inferred from mutant phenotype (Jiang et al., 2000)
Cellular Component
CV term Evidence References
nucleus inferred from direct assay (Ward et al., 2006)
Terms Based on Predictions or Assertions ( 10 terms )
Molecular Function
CV term Evidence References
sequence-specific DNA binding transcription factor activity non-traceable author statement (FlyBase, 1992-) traceable author statement (Gibson and Schubiger, 2001) zinc ion binding inferred from electronic annotation with InterPro:IPR007087, InterPro:IPR015880 (FlyBase Curators et al., 2004-)
Biological Process
CV term Evidence References
cell death traceable author statement (Thummel, 2001) ecdysone-mediated induction of salivary gland cell autophagic cell death non-traceable author statement (Buszczak and Segraves, 2000) metamorphosis non-traceable author statement (Swiss-Prot Project Members, 1995.6.1, Swiss-Prot Project Members, 2000.10.1) traceable author statement (Tissot and Stocker, 2000) regulation of development, heterochronic traceable author statement (Thummel, 2001) regulation of transcription, DNA-dependent non-traceable author statement (Swiss-Prot Project Members, 1995.6.1, Swiss-Prot Project Members, 2000.10.1) salivary gland cell autophagic cell death non-traceable author statement (Lee and Baehrecke, 2000) salivary gland histolysis non-traceable author statement (Farkas et al., 2002)
Cellular Component
CV term Evidence References
nucleus non-traceable author statement (Farkas et al., 2002, FlyBase, 1992-, Swiss-Prot Project Members, 1995.6.1, Swiss-Prot Project Members, 2000.10.1)
Sequence Ontology: Class of Gene
	nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
Protein-protein
Interacting group Assay References
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions br allele Gene References br1 bs (Ward et al., 2003) E(br)24 (Ward et al., 2003) E(br)65 (Ward et al., 2003) E(br)72 (Ward et al., 2003) E(br)121 (Ward et al., 2003) E(br)155 (Ward et al., 2003) E(br)160 (Ward et al., 2003) E(br)187 (Ward et al., 2003) E(br)333 (Ward et al., 2003) E(br)420 (Ward et al., 2003) E(br)444 (Ward et al., 2003) LIMK1 (Chen et al., 2004) Rho1 (Ward et al., 2003, Bayer et al., 2003, Chen et al., 2004) Sb (Ward et al., 2003) Sec61α (Ward et al., 2003) Tm1 (Ward et al., 2003) zip (Chen et al., 2004) br2Bc-1 B (Brennan et al., 2001) br2Bc-2 Eip74EF (Fletcher and Thummel, 1995) br5 B (Brennan et al., 2001) bs (Ward et al., 2003) E(br)65 (Ward et al., 2003) E(br)121 (Ward et al., 2003) E(br)333 (Ward et al., 2003) E(br)444 (Ward et al., 2003) Eip74EF (Fletcher and Thummel, 1995) Rho1 (Ward et al., 2003) Sb (Ward et al., 2003) brBRcore.TNT.Q1.Z1.hs ftz-f1 (Yamada et al., 2000) brnpr-3 B (Brennan et al., 2001) brrbp-1 Msex\br (Bayer et al., 2003) brrbp-5 B (Brennan et al., 2001) brZ2.hs ftz-f1 (Yamada et al., 2000) unspecified Mef2 (Lovato et al., 2005) Met (Wilson et al., 2006) Sb (Hammonds and Fristrom, 2006, Gotwals and Fristrom, 1991) twi (Lovato et al., 2005) zip (Gotwals and Fristrom, 1991)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 69091 DroID - A comprehensive database of gene and protein interactions. • FBgn0000210 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 44505
Orthologs
Genome-wide drosophilid orthologs	Dgri\GH17956 Dper\GL14266 Dpse\GA22724 Dvir\GJ16036 Dwil\GK18522
Curated drosophilid orthologs	Dfun\br   Dhyd\br   Dkan\br   Dmer\br   Dpar\br   Dpse\GA11031   Dpse\broad   Drep\br   Dvir\br
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AGAP002165-PA Anopheles gambiae (mosquito) XP_001942681.1 Acyrthosiphon pisum (pea aphid) 111317 Branchiostoma floridae (lancelet) ENSCAFP00000014684 Canis familiaris (dog) ENSDARP00000081229 Danio rerio (zebra fish) XP_001602030.1 Nasonia vitripennis (wasp) ENSORLP00000016699 Oryzias latipes (japanese killifish) PHUM005619-PA Pediculus humanus (human body louse) ENSPPYP00000004648 Pongo pygmaeus (orangutan) XP_974947.2 Tribolium castaneum (red flour beetle) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB14396 NV15002 LH21267 FBgn0220520 FBgn0203221 JGI_V11_39364 JGI_V11_59111 FBgn0244127 FBgn0151871 FBgn0139239 FBgn0125424 FBgn0105142 FBgn0099163 CPIJ003269 BGIBMGA009907 ACYPI008576 GB30150 GB14070 AGAP002165 AAEL008426 PHUM258420 GLEAN_05474
Stocks & Reagents
Stocks Listed in FlyBase ( 49 )
Bloomington	3994 br1 29972 y1 br5/Binsn 29969 y1 br2Bc-2/Binsn 36561 y1 brnpr-5/Binsn 36562 y1 brnpr-6/Binsn
Harvard	- P{XP}brd03598 - P{XP}brd06704 - PBac{PB}brc00809
Kyoto	107549 br1
VDRC	v104648 P{KK111704}VIE-260B v13705 w1118; P{GD4279}v13705 v38526 w1118; P{GD7318}v38526
	More stocks available...
Genomic Clones ( 1 )
	BACR33A08 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 127 )
	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	LP12157
Other clones	MIP22851
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	LP01363 LP05017 LP08009 LP10381 LP11337 LP15786 LP16818 LP17678 LP19054 LP19378 LP20362 LP23110 RH46193 RH49544
Other clones	5349 5778 7395 30745 30877 32068 33596 37838 40161 41501 41725 53926 58426 61103 64729 65936 67385 74507 78262 78578 82428 102367 144929 147475 172512 175191 175831 183448 191608 192732 214467.41 277163 282908 525084 CK01.8D.E8 CK01.10A.D3 CK01.16B.C2 CK01.28B.D8 CK01.35C.A10 CK01.35C.A12 CK01.91A.G3 CK01.92A.G3 CK01.96A.E10 CK01.107C.E6 CK01.113B.E1 CK01.124C.G2 CK01.127A.D5 EC14543 EC22020 EC33962 EK011452 EK016131 EK030626 EK039150 EK039923 EK071326 EK071926 EK074749 EK090451 EK094657 EK097315 EK102001 EK124445 EK125641 EK142425 EK150015 EK153951 EK154612 EK166326 EK166529 EK178061 EK187129 EK189905 EK193047 EK196035 EK197502 EK203007 EK207530 EK215555 EK219407 EK247406 EK250957 EK276316 EK280326 EK280638 EK288533 EK291058 EK294160 EK295640 EK298746 EK298830 EK304442 EK304908 EK308438 EP05519 EP06501 EP08618 EP10539 EP13357 EP21745 ESG014d.E21_A04 ESG018b.E21.1_A07 ESG0112a.B21.1_E01 LP10481 RH09228 RP004170439 RP004175700 RP004210357 RP004339478 RP004377828 RP004420091
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0000210 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 44505
Antibody Information
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of	Source for merge of: br CG11491 (Bayraktaroglu, 2000.4.12) Source for merge of: br CG11511 (FlyBase Genome Annotators, 2002-2003) Source for merge of: br CG11514 (Ashburner, 2000.5.29) Source for merge of: br l(1)G0318 l(1)G0401 l(1)G0018 l(1)G0042 l(1)G0284a (Peter et al., 2002)
Additional comments	Annotations CG11491, CG11514 and CG11511 merged as CG11491 in release 3 of the genome annotation. (FlyBase Genome Annotators, 2002-2003) 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
	The br product prevents rho expression in egg chambers. (Ward et al., 2006) br appears to be required for Ddc activation eclosion. (Chen et al., 2002) The ecdysone pathway is required for proliferation and differentiation of hematopoitic precurssors of lamellocytes and crystal cells. (Sorrentino et al., 2002) br isoforms regulate morphogenetic furrow progression and photoreceptor specification in the developing eye. (Brennan et al., 2001) br is required for salivary gland, autophagic, cell death. (Lee and Baehrecke, 2001) rbp+ function is required in the dorsal epidermal muscle attachment site for the correct attachment of the indirect flight muscles. (Sandstrom and Restifo, 1999) The target genes of the BR-C (br) in oogenesis include a protein essential for endoreplication and chorion gene amplification. (Tzolovsky et al., 1999) 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. (Bayer et al., 1997) br is expressed in a bilaterally symmetrical pattern in the lateral-dorsal-anterior follicle cells during late oogenesis. (Deng and Bownes, 1997) br is essential for the morphogenesis of dorsal appendages. (Deng and Bownes, 1997) 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. (Sandstrom et al., 1997) 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. (Bayer et al., 1996) 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. (Crossgrove et al., 1996) br gene participates in parallel tissue-specific ecdysone response cascades to Hsp23. (Dubrovsky et al., 1996) 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'. (Bayer et al., 1995) 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. (Bayer et al., 1995) 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. (Fletcher and Thummel, 1995) Hormonal induction of Ddc in the epidermis is mediated by br. (Hodgetts et al., 1995) 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. (Restifo et al., 1995) 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. (Dubrovsky et al., 1994) Differential expression of Broad complex transcription factors may forecast tissue-specific developmental fates during metamorphosis. (Emery et al., 1994) br function is required for the proper timing of Sgs4 induction and acts through sequences located outside the Sgs4-Pig1 intergenic region. (von Kalm et al., 1994) Ecdysteroid-regulated gene. (Andres et al., 1993) 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. (Karim et al., 1993) Mosaic experiments reveal a somatic but not germ line requirement for npr function. (Huang and Orr, 1992) Developmentally regulated peak of arginine kinase at pupal stage P3 is affected by altering copy number of Eip75B and BRC, but not Eip74EF. (James and Collier, 1992) The temporal pattern of br transcription during the third larval instar stage has been analysed. (Karim and Thummel, 1992) 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. (Kokoza et al., 1992) Mutations at br disrupt the metamorphosis of internal tissues. (Restifo and White, 1992) When the 2B region is subjected to position effect the br locus in inactivated. (Belyaeva and Zhimulev, 1991) The mlf phenotype (malformed syndrome), wing malformations and leg defects, can be produced by a dominant genetic interaction between Sb and br. (Gotwals and Fristrom, 1991) '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. (Guay and Guild, 1991) 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. (Hofmann et al., 1991) 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. (Kokoza et al., 1991) 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. (Mazina and Korochkina, 1991) 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. (Nelson et al., 1991) Mutations at br reduce the transcription rate or stability of the small heat shock protein mRNAs. (Dubrovsky et al., 1990) Female sterile mutations at the br locus do not interfere with the expression of Cp70. (Mazina and Dubrovskii, 1990) 'l(1)2Bc' is involved in imaginal disc fusion. br is involved in imaginal disc evagination. (Sampedro et al., 1989) 'br' and 'l(1)2Bc' alleles are fully complementing. (Kiss et al., 1988) 'l(1)2Bc' function is involved in appendage elongation. (Kiss et al., 1988) 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.
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_080020 NM_166893 NM_166894 NM_166895 NM_166896 NM_166897 NM_166898 NM_001144672 NM_001169167 NM_001201596 NM_001201597 NM_001201598 RefSeq (Proteins) • NP_524759 NP_726749 NP_726750 NP_726751 NP_726752 NP_726753 NP_726754 NP_001138144 NP_001162638 NP_001188525 NP_001188526 NP_001188527 DDBJ / EMBL / GenBank • AAB09760 AAF45647 AAF45648 AAF45650 AAF45651 AAL39901 AAN09052 AAN09053 AAN09054 ACL82877 ACZ95175 ADV37609 ADV37610 ADV37611 AF345689 AF345690 AF345691 AF345692 AF345693 AF345694 AF345695 AF345696 AF345697 AF345698 AF345699 AF345700 AF345701 AI260963 AI296538 AI297782 AJ300017 AJ300018 AJ300019 AJ300020 AJ300021 AL009146 AL021067 AW944129 AW944154 AY069756 BZ748779 CAA15626 CAA15627 CAA15628 CAA15629 CAA38474 CAA38475 CAA38476 CAA38477 CZ472859 CZ473454 CZ475289 CZ475290 CZ475657 CZ476127 DQ327739 S55610 S55612 U51585 X54663 X54664 X54665 X54666 X62380 X63618 X97997 X97998 Z32013 Z32190 Z32387 Z32433 Entrez Gene - A searchable database of RefSeq genes. • 44505 UniProtKB/Swiss-Prot • Q01295 Q24206 UniProtKB/TrEMBL • E1JJB2 Q8IRX4 Q9W572
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • BTB/POZ-like (IPR000210) Zinc finger, C2H2 (IPR007087) BTB/POZ (IPR013069) BTB/POZ fold (IPR011333) Zinc finger, C2H2-like (IPR015880)
Linkouts
	BioGRID - A database of protein and genetic interactions • 69091 DroID - A comprehensive database of gene and protein interactions. • FBgn0000210 DRSC - Results from RNAi screens. • FBgn0000210 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0000210 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG11491-RC;CG11491-RA;CG11491-RF;CG11491-RD;CG11491-RE FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0000210 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 44505 InParanoid A subset of ortholog calls from InParanoid. • AGAP002165-PA Anopheles gambiae (mosquito) XP_001942681.1 Acyrthosiphon pisum (pea aphid) 111317 Branchiostoma floridae (lancelet) ENSCAFP00000014684 Canis familiaris (dog) ENSDARP00000081229 Danio rerio (zebra fish) XP_001602030.1 Nasonia vitripennis (wasp) ENSORLP00000016699 Oryzias latipes (japanese killifish) PHUM005619-PA Pediculus humanus (human body louse) ENSPPYP00000004648 Pongo pygmaeus (orangutan) XP_974947.2 Tribolium castaneum (red flour beetle) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /newgene/broadcx1.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 44505 modMine - Data generated by the modENCODE project. • FBgn0000210 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB14396 NV15002 LH21267 FBgn0220520 FBgn0203221 JGI_V11_39364 JGI_V11_59111 FBgn0244127 FBgn0151871 FBgn0139239 FBgn0125424 FBgn0105142 FBgn0099163 CPIJ003269 BGIBMGA009907 ACYPI008576 GB30150 GB14070 AGAP002165 AAEL008426 PHUM258420 GLEAN_05474 REDfly - A database of transcriptional regulatory elements. • FBgn0000210
Synonyms & Secondary IDs ( 97 )
Reported As
Symbol Synonym	2B5 (Cox and Collier, 1998, Truman et al., 1993) 2Bc (Lee et al., 2002, von Kalm et al., 1994, Basso et al., 2006, Consoulas et al., 2005, Spokony and Restifo, 2009) A18 (Kramers et al., 1991, Kramers et al., 1983) br (Khokhar et al., 2008, Hammonds and Fristrom, 2006, Zhang et al., 2007, Sorrentino et al., 2002, Alekseyenko et al., 2006, Riddiford et al., 2003, Chen and Schupbach, 2006, Zhou and Riddiford, 2008, Aerts et al., 2007, Zhou et al., 2007, Cook et al., 2008.9.3, Minakuchi et al., 2008, Basso et al., 2006, Wilson et al., 2006, Lembong et al., 2008, Wang et al., 2008, Pearse et al., 2008, Zhang et al., 2008, Weake et al., 2008, Grieder et al., 2007, Zeitouni et al., 2007, Christensen et al., 2008.12.28, Yakoby et al., 2008, Yakoby et al., 2008, Gilchrist et al., 2008, Stofanko et al., 2008, Weake et al., 2011, Zartman et al., 2008, Lembong et al., 2009, Bernardi et al., 2009, Mirth et al., 2009, Dworkin et al., 2009, Zhou et al., 2004, Liu and Lehmann, 2008, Zartman et al., 2009, Perkins et al., 2009.8.10, Cui et al., 2009, Manak et al., 2006, Consoulas et al., 2005, Zhou et al., 2009, Parrish et al., 2009, Dalton et al., 2009, Cao et al., 2007, Kim et al., 2010, Bai et al., 2009, Xiang et al., 2010, Klebes et al., 2002, Spokony and Restifo, 2009, Seong et al., 2011) BR (Zhou and Riddiford, 2003, Boyle and Berg, 2008, Boyle and Berg, 2009) Br (Yakoby et al., 2007, Altman and Truman, 2007, Herranz et al., 2010, Rewitz et al., 2010) BR_C (Tissot and Stocker, 2000) BR-C (Cakouros et al., 2008, Kelly and Daniel, 2006, Badenhorst et al., 2005, Caceres and Nilson, 2005, Kilpatrick et al., 2005, Farkas et al., 2005, Terashima and Bownes, 2004, Martin and Baehrecke, 2004, Dubrovsky et al., 2004, Chiang and Kurnit, 2003, Mazo et al., 2004, Nakamura and Matsuno, 2003, James and Berg, 2003, Pokholkova and Zhimulev, 2002, Cakouros et al., 2002, Zhang et al., 2002, Kumar et al., 2003, Bialecki et al., 2002, Gonzy et al., 2002, Kozlova and Thummel, 2000, Lafont, 2000, Duncan and Warrior, 2002, Thummel and Chory, 2002, Lee et al., 2002, Gonzalez et al., 2002, Zhou and Riddiford, 2001, Lee et al., 2002, Zhou and Riffiford, 2002, Berger et al., 2002, Simon et al., 2001, Lee et al., 2001, Deng et al., 2001, Dubrovsky et al., 2001, Suzanne et al., 2001, Farkas and Mechler, 2001, Lee et al., 2001, Thummel, 2001, Beckstead et al., 2001, Hsu and Schulz, 2000, Lee and Baehrecke, 2001, Buszczak and Segraves, 2000, Baehrecke, 2000, Yamada et al., 2000, Lee et al., 2000, Li and Bender, 2000, Jiang et al., 2000, Lee and Baehrecke, 2000, Peri and Roth, 2000, Bayer, 1996.3.16, Zhao and Bownes, 1999, Buszczak et al., 1999, Lam et al., 1999, Roy and VijayRaghavan, 1999, Broadus et al., 1999, Roth et al., 1999, Baehrecke and Lee, 1999, Hall and Thummel, 1998, Buszczak and Segraves, 1998, Deng and Bownes, 1998, Brennan et al., 1998, Doronkin and Korochkin, 1998, D'Avino and Thummel, 1998, Fisk and Thummel, 1998, Bender et al., 1997, Thummel, 1997, Bayer et al., 1997, Atkinson, 1994, Lam et al., 1997, Makunin et al., 1996, Ito et al., 1996, Baehrecke, 1996, D'Avino et al., 1995, Huet et al., 1993, Restifo et al., 1992, Karim and Thummel, 1992, Galceran et al., 1990, Farkas et al., 2006, Atkey et al., 2006, Choi et al., 2006, Legube et al., 2006, Hoopfer et al., 2008, Furuhashi et al., 2006, Daish et al., 2003, Rendina et al., 2010, Thomas and Cripps, 2008, Jindra and Konopova, 2008, Thummel, 2007, Ciurciu et al., 2006, Bashirullah et al., 2007, Peretz et al., 2007, Shravage et al., 2007, McBrayer et al., 2007, Cao et al., 2007, Kozlova et al., 2009, Chittaranjan et al., 2009, Bernardo et al., 2009, Lachance et al., 2009, Beckstead et al., 2005, Sawatsubashi et al., 2010, Ritter and Beckstead, 2010, Lin et al., 2011, Francis et al., 2010, Dialynas et al., 2010) BRC (Mendjan et al., 2006, Yerushalmi et al., 2002, Krempler and Brenig, 1999, Restifo and Wilson, 1998, Liu et al., 1996, Bayer et al., 1996, Jindra, 1994, Andres et al., 1993, Spokony and Restifo, 2007, Costantino et al., 2008, Consoulas et al., 2005, Restifo and Hauglum, 1998, Spokony and Restifo, 2009) BR-c (Chen et al., 2002) br-C (Bourbon et al., 2002) Br-C (Semeshin et al., 2001, Ghbeish et al., 2000, Henrich and Brown, 1995, Chen and Schupbach, 2006, Papadia et al., 2005, Maurange et al., 2008, Kalifa et al., 2009, Semeshin et al., 2002) BR-C Z1 (Farkas et al., 2003, Sedkov et al., 2003, Farkas et al., 2002, Dequier et al., 2001) BrC-Z1 (Ghbeish and McKeown, 2002, Ghbeish et al., 2001) BRC-Z1 (Schubiger and Truman, 2000, Lovato et al., 2005, Costantino et al., 2008) BRC-Z2 (Costantino et al., 2008) Br-C Z2 (Terashima and Bownes, 2006) BRC-Z3 (Costantino et al., 2008, Consoulas et al., 2005) Br-C-Z3 (Terashima and Bownes, 2006) BRC-Z4 (Costantino et al., 2008) br-Z1 (Schubiger and Truman, 2004, Rifkin et al., 2003) br-Z3 (Rifkin et al., 2003) Br-Z4 (Bauer et al., 2010) CG11491 (Khokhar et al., 2008, Bourbon et al., 2002, Benos, 2000.12.13, Alekseyenko et al., 2006, Grieder et al., 2007, Keleman et al., 2009.8.5, Ni et al., 2010.12.1) CG11511 (Benos, 2000.12.13, Ashburner, 2000.5.29, ) CG11514 (Benos, 2000.12.13, Ashburner, 2000.5.29, ) de12 (Orr et al., 1989) ecs (Weiler and Wakimoto, 1995, Zhimulev et al., 1995, Solov'eva, 1992, Segal, 1993, Balasov and Bgatov, 1992, Kokoza et al., 1992, Kokoza et al., 1991, Protopopov et al., 1991, Umbetova et al., 1991, Belyaeva and Zhimulev, 1991, Dubrovsky et al., 1990, Mazina and Dubrovskii, 1990, Mazina et al., 1990, Solovyeva and Belyaeva, 1989, Demakova and Belyaeva, 1988, Belyaeva et al., 1987, ) EG:17A9.1 (Benos et al., 2001, Benos, 2000.12.13, ) EG:25D2.1 (Benos et al., 2001, ) EG:123F11.1 (Benos et al., 2001, ) l(1)2Ba   l(1)2Bab (Dubrovsky et al., 1994, Merriam et al., 1991, Thummel, 1990, ) l(1)2Bad   l(1)2Bb   l(1)2Bc (Dubrovsky et al., 1994, Merriam et al., 1991, Dubrovsky et al., 1990, Thummel, 1990, ) l(1)2Bd (Merriam et al., 1991, Thummel, 1990, ) l(1)d.norm.1   l(1)dn1   l(1)d norm-12 (Gatti and Baker, 1989) l(1)ESHS5 (Merriam et al., 1991, ) l(1)G0018 (Schaefer et al., 1999.3.29, ) l(1)G0042 (Schaefer et al., 1999.8.12, ) l(1)G0284   l(1)G0284a (Benos et al., 2001, Schaefer et al., 1999.6.15, ) l(1)G0318 (Schaefer et al., 1999.6.15, ) l(1)G0401 (Schaefer et al., 1999.8.12, ) l(1)n34   l(1)npr1 (Segal, 1993, ) l(1)npr-1 (Hofmann et al., 1991, Martin et al., 1989, Vijay Raghavan et al., 1988) l(1)PP1 (Solov'eva, 1992, Mazina et al., 1991, Mazina et al., 1990) l(1)pp-1 (Demakova et al., 1988, Belyaeva et al., 1980) l(1)pp1 (Aizenzon and Belyaeva, 1982, Belyaeva et al., 1982, ) l(1)pp-2 (Demakova et al., 1988, Belyaeva et al., 1980) l(1)pp2 (Belyaeva et al., 1982, ) l(1)ts132   l(1)ts144   l(1)ts358   l(1)ts376   npr (Dunne et al., 2002, Zhou et al., 1999, Lehmann and Korge, 1996, Dubrovsky et al., 1994, ) npr1 (Crossgrove et al., 1996, Baehrecke and Thummel, 1995, von Kalm et al., 1994, Restifo and Merrill, 1994, Restifo and White, 1992, Merriam et al., 1991, Gotwals and Fristrom, 1991, Restifo and White, 1991, Thummel, 1990, Kiss et al., 1988, Basso et al., 2006, Consoulas et al., 2005, Spokony and Restifo, 2009) npr-1 (Galceran et al., 1990) nprl (Hodgetts et al., 1995) nrp (Dunne et al., 2002) nrp1 (Consoulas et al., 2005) o.c.c.   PP1 (Lioultcheva et al., 2002, Alphey et al., 2000, Belyaeva et al., 1987) PP2 (Belyaeva et al., 1987) rbp (Martin and Baehrecke, 2004, von Kalm et al., 2003, Farkas et al., 2002, Wilson et al., 2002, Restifo et al., 1997, Fletcher et al., 1997, Crossgrove et al., 1996, Riddiford, 1993, Thummel, 1990, Basso et al., 2006, Consoulas et al., 2005, Spokony and Restifo, 2009) rdp   rds (Merriam et al., 1991, ) unnamed (Georgel et al., 1991, Kiss et al., 1978) uq (Fahmy and Fahmy, 1958, ) Z1 (Restifo et al., 1993, Basso et al., 2006) Z2 (Restifo et al., 1993, Basso et al., 2006) Z3 (Restifo et al., 1993, Basso et al., 2006) Z4 (Gortchakov et al., 2004, Restifo et al., 1993, Eggert et al., 2005, Basso et al., 2006)
Name Synonym	broad (Khokhar et al., 2008, Huang, 1993.4.13, Huang, 1993.4.13, Zhang et al., 2007, Terry et al., 2006, Barmina et al., 2005, Zhou and Riddiford, 2008, Zhou et al., 2007, Minakuchi et al., 2008, Ward et al., 2006, Wilson et al., 2006, Lembong et al., 2008, Pearse et al., 2008, Grieder et al., 2007, Zeitouni et al., 2007, Stofanko et al., 2008, Bernardi et al., 2009, Mirth et al., 2009, Zhou et al., 2004, Schaaf et al., 2009, Consoulas et al., 2005, Zhou et al., 2009, Parrish et al., 2009, Dalton et al., 2009, Patch et al., 2009, Kim et al., 2010, Bai et al., 2009, Xiang et al., 2010, Sharma et al., 2011, Klebes et al., 2002) Broad (Wahlstrom et al., 2006, Siddall et al., 2006, Cho et al., 2005, Jordan et al., 2005, Riddiford et al., 2003, Adryan and Teichmann, 2007, Boyle and Berg, 2008, Yakoby et al., 2008, Zartman et al., 2008, Cui et al., 2009, Boyle and Berg, 2009, Truman et al., 2010) broad complex (Lehmann, 2004, Gibson and Schubiger, 2001) broad-complex (Cho-Fertikh et al., 2003) Broad-Complex (Guichet et al., 2001, Makunin, 1996.5.21, Llanos, 1991.12.17, Kholodilov, 1996.5.21, Kozlova and Thummel, 2000, Jindra and Konopova, 2008, Papadia et al., 2005, Thummel, 2007, Kalifa et al., 2009) Broad Complex (Consoulas et al., 2001, Llanos, 1991.12.23, Tissot and Stocker, 2000, Maines et al., 2004, Farkas and Mechler, 2001, Lovato et al., 2005, Spokony and Restifo, 2007, Vielmas et al., 2008, Costantino et al., 2008, Maurange et al., 2008, Consoulas et al., 2005, Lachance et al., 2009, Restifo and Hauglum, 1998, Spokony and Restifo, 2009) Broad-complex (Buszczak and Segraves, 2000, Roy and VijayRaghavan, 1999, ) Broad complex (Thomas and Cripps, 2008) 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 (Fahmy and Fahmy, 1958, )
Secondary FlyBase IDs
	FBgn0000801 FBgn0001373 FBgn0001374 FBgn0001375 FBgn0001644 FBgn0001684 FBgn0002964 FBgn0003222 FBgn0005389 FBgn0005581 FBgn0010011 FBgn0026711 FBgn0027219 FBgn0027234 FBgn0028290 FBgn0028364 FBgn0029583 FBgn0029585 FBgn0029586
References ( 496 )
Generate a list of	All references relating to this gene ( 496 )
List References by type	Research paper  ( 301 ) Supplementary material  ( 8 ) Review  ( 49 ) Personal communication to FlyBase  ( 17 ) Abstract  ( 94 ) FlyBase analysis  ( 4 ) DNA/RNA sequence record  ( 7 ) Letter  ( 1 ) Protein sequence record  ( 2 ) Book  ( 4 ) Computer file  ( 4 ) Curated genome annotation  ( 2 ) Automatic genome annotation  ( 1 ) Book review  ( 1 ) Stock list  ( 1 )
Recent research papers ( 18 )
	Gradilla et al., 2011, Genetics 189(3): 871--883 Isoform-Specific Regulation of a Steroid Hormone Nuclear Receptor by an E3 Ubiquitin Ligase in Drosophila melanogaster. [FBrf0216551] Huang et al., 2011, Development 138(11): 2283--2291 DPP-mediated TGF{beta} signaling regulates juvenile hormone biosynthesis by activating the expression of juvenile hormone acid methyltransferase. [FBrf0213687] Lin et al., 2011, Insect Mol. Biol. 20(1): 87--95 Hormone receptor-like in 96 and Broad-Complex modulate phenobarbital induced transcription of cytochrome P450 CYP6D1 in Drosophila S2 cells. [FBrf0212740] Seong et al., 2011, Biogerontology 12(2): 93--107 Genome-wide analysis of low-dose irradiated male Drosophila melanogaster with extended longevity. [FBrf0213307] Sharma et al., 2011, Chemosphere 82(3): 370--376 Transcriptome analysis provides insights for understanding the adverse effects of endosulfan in Drosophila melanogaster. [FBrf0212610] Weake et al., 2011, Genes Dev. 25(14): 1499--1509 Post-transcription initiation function of the ubiquitous SAGA complex in tissue-specific gene activation. [FBrf0214372] Zartman et al., 2011, Phys. Biol. 8(4): 045003 Pattern formation by a moving morphogen source. [FBrf0214496] Bauer et al., 2010, BMC Bioinformatics 11: 366 Dual-functioning transcription factors in the developmental gene network of Drosophila melanogaster. [FBrf0211408] Blanco et al., 2010, BMC Dev. Biol. 10: 94 Gene expression following induction of regeneration in Drosophila wing imaginal discs. Expression profile of regenerating wing discs. [FBrf0211812] Dialynas et al., 2010, Development 137(18): 3067--3077 The role of Drosophila Lamin C in muscle function and gene expression. [FBrf0211601] Francis et al., 2010, Curr. Biol. 20(20): 1799--1808 dDOR Is an EcR Coactivator that Forms a Feed-Forward Loop Connecting Insulin and Ecdysone Signaling. [FBrf0212109] Herranz et al., 2010, Mol. Ecol. 19(19): 4255--4264 Spaceflight-related suboptimal conditions can accentuate the altered gravity response of Drosophila transcriptome. [FBrf0216346] Kim et al., 2010, BMC Cell Biol. 11: 9 A genetic screen for modifiers of Drosophila caspase Dcp-1 reveals caspase involvement in autophagy and novel caspase-related genes. [FBrf0209948] Rewitz et al., 2010, Dev. Cell 19(6): 895--902 Steroid Hormone Inactivation Is Required during the Juvenile-Adult Transition in Drosophila. [FBrf0212489] Ritter and Beckstead, 2010, Dev. Dyn. 239(10): 2685--2694 Sox14 is required for transcriptional and developmental responses to 20-hydroxyecdysone at the onset of drosophila metamorphosis. [FBrf0212221] Sawatsubashi et al., 2010, Genes Dev. 24(2): 159--170 A histone chaperone, DEK, transcriptionally coactivates a nuclear receptor. [FBrf0209754] Truman et al., 2010, Development 137(1): 53--61 Role of Notch signaling in establishing the hemilineages of secondary neurons in Drosophila melanogaster. [FBrf0209606] Xiang et al., 2010, Dev. Biol. 344(2): 800--808 br regulates the expression of the ecdysone biosynthesis gene npc1. [FBrf0211381] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...