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Gene Dmel\bw

General Information
Symbol	Dmel\bw	Species	D. melanogaster
Name	brown	Annotation symbol	CG17632
Feature type	protein_coding_gene	FlyBase ID	FBgn0000241
Gene Model Status	Current	Stock availability	2369 publicly available
Genomic Location
Chromosome (arm)	2R	Recombination map	2-104.5
Cytogenetic map	59E2-59E3	Sequence location	2R:19,415,328..19,426,016 [-]
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 brown is referred to in FlyBase by the symbol Dmel\bw (CG17632, FBgn0000241). It is a protein_coding_gene from Drosophila melanogaster. Based on sequence similarity, it is predicted to have molecular function: ATPase activity, coupled to transmembrane movement of substances. There is experimental evidence that it is involved in the biological process: ommochrome biosynthetic process. 156 alleles are reported. The phenotypes of these alleles are annotated with: ocellus; hemolymph; eye; wing; pigment granule; pigment cell. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: ABC transporter, conserved site; ABC transporter-like; ABC-2 type transporter; ATPase, AAA+ type, core; Pigment precursor permease. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderate expression to a trough of very low expression. Peak expression observed within 06-18 hour embryonic stages, during late larval stages, at stages throughout the pupal period, in adult male 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 Malpighian tubules. Expression at moderate levels in the following post-embryonic organs or tissues: adult eye, adult Malpighian tubules, larval fat body. Gene sequence location is 2R:19415328..19426016.
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	bw: brown Eye color light brownish wine on emergence, darkening to garnet. Red pigments lacking. Xanthommatin mostly replaced by dihydroxanthommatin (Phillips, Forrest, and Kulkarni, 1973, Genetics 73: 45-56). Pigment granules present but somewhat smaller than in wild type. Adult testes and vasa colorless. Larval Malpighian tubules pale yellow (Beadle, 1937, Genetics 22: 587-611). Produces white eyes in combination with v, cn, or st. Eye color autonomous when transplanted into wild-type host (Beadle and Ephrussi, 1936, Genetics 21: 230). RK1. *bwA: brown-Auburn Nearly uniform brown but with extra Y chromosome shows strong variegation. Homozygote usually lethal. RK1A. bw4 Homozygotes have normal eye color; in heterozygotes with other alleles, red pigment is reduced. bw4/bw5 is purpleoidlike. bw4/bw like bw but darker. RK3. bw5 bw5/bw4 is purpleoidlike (see bw4); bw5/bw is light yellowish brown; bw5/+ is wild type; bw5/bw5 is lethal. RK2A. bw6 A subliminal recessive allele of bw; homozygotess are wild type in all combinations except in flies that are homozygous or hemizygous for wco2, which are like bw or wco. bw6/bw4 suppresses wco2, but all other heterozygotes between bw6 and bw alleles which were tested did not suppress. RK1 in combination with wco2. bwa: brown-amber Eye color light brownish yellow. Adult testes and vasa colorless. Larval Malpighian tubules slightly paler yellow than wild type. bwa/bw gives eye color slightly lighter than bw. RK1. bwD: brown-Dominant Eye color varies with age from purple to brown. Shows slight variegation in combination with st (Slatis, 1955). Wings pebbled. bwD/+ shows nearly a 100-fold reduction in pteridine levels; no accumulation of bw RNA detectable. bwD/bwVI > bwD/+ > bwD/bwD in severity of effect (Henikoff, and Dreesen, 1989, Proc. Nat. Acad. Sci. USA 86: 6704-08). Variegation suppressed by extra Y chromosomes (Brosseau, 1959, DIS 33: 123). Homozygote viable and fertile. Larval Malpighian tubules bright yellow (Brehme and Demerec, 1942, Growth 6: 351-56). RK1A. *bwptm: brown:pteridine modifier Undetectable in wild-type background. In wsat or wcf flies that are at the same time v, cn, or st, the amount of red eye pigment in +/+ > +/bwptm > bwptm/bwptm. bwR: brown-Rearranged These rearranged derivatives exhibit variegated phenotype in combination with +. Mostly homozygous lethal; survivors have brown eyes. bwV1: brown-Variegated Eye color like bw or pr, mottled with darker spots that deepen in red color with age. With st or v, has pale orange ground with dark orange spots. bwVI/bw shows sharply reduced levels of transcript from both the bw+ allele in cis with the rearrangement and the bw allele on the homologous second chromosome (Henikoff, and Dreesen, 1989, Proc. Nat. Acad. Sci. USA 86: 6704-08). Extra Y chromosome, as with other variegated browns, suppresses brown color, giving red eye sparsely speckled or splotched with darker spots. Amount of wild-type eye color also responds to fourth-chromosome constitution; 4/C(4) > C(4) = 4/4 > 4/0, with the first approaching wild type and the latter nearly bw (Lindsley and Rokop). Larval Malpighian tubules normal (Glass, Brehme). Generally lethal homozygous and in combination with other brown-Variegateds. Heterozygotes fully viable and fertile. RK1A.
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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FB2012_01	Sequence features HFA04005 HFA04677 BKN24865
FB2011_10
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 59E2-59E3   Limits computationally determined from genome sequence between P{lacW}l(2)s4830s4830&P{lacW}chrwk06908 and P{PZ}Dcp-102132
Experimentally Determined Cytological Location
Cytogenetic map Notes References 59E1-59E2     (Dreesen et al., 1988)
Experimentally Determined Recombination Data
Location	2-104.5 (Nitasaka et al., 1995, Fernandez Gianotti, 1950) 2-105 (Farmer, 1974)
Left of (cM)	or Kr sp (Ives, 1967)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\bw for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
	Known mutation in sequenced strain: premature stop Transposon inserted in intron
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) bw-RA FBtr0072117  NM_079100   2732   675 bw-RB FBtr0114457  NM_206206   1256   328
Additional Transcript Data & Comments
Reported size (kB)	3.0, 2.8 (northern blot) (Dreesen et al., 1988)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein bw-PA   FBpp0072026   75.9   675   7.41     NP_523824   AAF47020 bw-PB   FBpp0112949   36.4   328   8.77     NP_995928   AAS64761
Additional Polypeptide Data & Comments
Reported size (kDa)	675 (aa) (Dreesen et al., 1988)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • ABC transporter-like (IPR003439) ATPase, AAA+ type, core (IPR003593) ABC-2 type transporter (IPR013525) Pigment precursor permease (IPR005284) ABC transporter, conserved site (IPR017871)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA140795   AA141233   AA141237   AA141417   AA141619   AA141698   AA141745   AA141987   AA142125   AA695666   AA696090   AA802073   AE013599 AAF47020   AE013599 AAS64761   AF047460   AI109818   AI134596   AI296359   AI532614   AI542319   AI544411   AW940450   AY095181   AY119503 AAM50157   BG640013   BI230577   BI232370   BI239344   BI239355   BI361306   BI364844   BI366326   BI592783   BT023281   BT082027   CK136030   CO176759   CO176870   CO179227   CO190457   CO190927   CO196145   CO196191   CO261856   CO262585   CO262614   CO262846   CO263114   CO263490   CO266285   CO266529   CO267517   CO268181   CO270415   CO271834   CO273139   CO273355   CO274492   CO274565   CO276008   CO276601   CO276918   CO278062   CO278614   CO278914   CO279906   CO280735   CO281510   CO283847   CO284272   CO284667   CO285073   CO285477   CO287895   CO290053   CO290439   CO292108   CO292886   CO296225   CO302531   CO302565   CO303725   CO304160   CO305150   CO305783   CO307366   CO309960   CO311832   CO314146   CO315482   CO315500   CO316498   CO316831   CO316845   CO317857   CO318413   CO318495   CO318549   CO319622   CO320245   CO320928   CO323471   CO323778   CO327476   CO328391   CO331098   CO331410   CO331946   CO331984   CO333255   CO333496   CO336449   CO338463   CO340064   CO340329   EC084370   EC198618   EC220947   EC226015   EC242196   EC247149   EC250127   EC254209   EL875429   GH943398   GH943771   GH944901   GH945458   GH947266   GH949771   GH951403   GH953628   GH956798   GH958199   GH959511   GH959530   GH960430   GH961590   GH966175   GH967286   GH967329   GH968662   GH971183   GH980251   GH980814   L05635 AAA28398   L23543 AAC37214   M20630 AAA28397   Z50313
UniProtKB/Swiss-Prot	P12428
UniProtKB/TrEMBL
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
Additional Descriptive Data
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
Additional Descriptive Data
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 FBgn0000241    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0000241 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 moderate expression to a trough of very low expression. Peak expression observed within 06-18 hour embryonic stages, during late larval stages, at stages throughout the pupal period, in adult male 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 FBgn0000241 expression level Developmental Stage   Expression Level embryo 00-02hr    39 embryo 02-04hr    218 embryo 04-06hr    312 embryo 06-08hr    791 embryo 08-10hr    393 embryo 10-12hr    493 embryo 12-14hr    450 embryo 14-16hr    553 embryo 16-18hr    282 embryo 18-20hr    111 embryo 20-22hr    65 embryo 22-24hr    52 larva L1    104 larva L2    80 larva L3 12hr old    36 larva L3 puffstage 1-2    76 larva L3 puffstage 3-6    446 larva L3 puffstage 7-9    905 white prepupae new    755 white prepupae 12hr    233 white prepupae 24hr    261 pupae 2d postWPP    956 pupae 3d postWPP    1191 pupae 4d postWPP    378 adult male 01day    362 adult male 05day    491 adult male 30day    487 adult female 01day    280 adult female 05day    111 adult female 30day    118 Expression Level Scale  None   Extremely low   Very low   Low   Moderate  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    39 embryo 02-04hr  (218) embryo 04-06hr  (312) embryo 06-08hr  (791) embryo 08-10hr  (393) embryo 10-12hr  (493) embryo 12-14hr  (450) embryo 14-16hr  (553) embryo 16-18hr  (282) embryo 18-20hr  (111) embryo 20-22hr    65 embryo 22-24hr    52 larva L1  104 larva L2    80 larva L3 12hr old    36 larva L3 puffstage 1-2    76 larva L3 puffstage 3-6  (446) larva L3 puffstage 7-9  (905) white prepupae new  (755) white prepupae 12hr  (233) white prepupae 24hr  (261) pupae 2d postWPP  (956) pupae 3d postWPP  (1191) pupae 4d postWPP  (378) adult male 01day  (362) adult male 05day  (491) adult male 30day  (487) adult female 01day  (280) adult female 05day  (111) adult female 30day  (118) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    39 embryo 02-04hr    218 embryo 04-06hr    312 embryo 06-08hr    791 embryo 08-10hr    393 embryo 10-12hr    493 embryo 12-14hr    450 embryo 14-16hr    553 embryo 16-18hr    282 embryo 18-20hr    111 embryo 20-22hr    65 embryo 22-24hr    52 larva L1    104 larva L2    80 larva L3 12hr old    36 larva L3 puffstage 1-2    76 larva L3 puffstage 3-6    446 larva L3 puffstage 7-9    905 white prepupae new    755 white prepupae 12hr    233 white prepupae 24hr    261 pupae 2d postWPP    956 pupae 3d postWPP    1191 pupae 4d postWPP    378 adult male 01day    362 adult male 05day    491 adult male 30day    487 adult female 01day    280 adult female 05day    111 adult female 30day    118 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    39 embryo 02-04hr    218 embryo 04-06hr    312 embryo 06-08hr    791 embryo 08-10hr    393 embryo 10-12hr    493 embryo 12-14hr    450 embryo 14-16hr    553 embryo 16-18hr    282 embryo 18-20hr    111 embryo 20-22hr    65 embryo 22-24hr    52 larva L1    104 larva L2    80 larva L3 12hr old    36 larva L3 puffstage 1-2    76 larva L3 puffstage 3-6    446 larva L3 puffstage 7-9    905 white prepupae new    755 white prepupae 12hr    233 white prepupae 24hr    261 pupae 2d postWPP    956 pupae 3d postWPP    1191 pupae 4d postWPP    378 adult male 01day    362 adult male 05day    491 adult male 30day    487 adult female 01day    280 adult female 05day    111 adult female 30day    118 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    39 embryo 02-04hr    218 embryo 04-06hr    312 embryo 06-08hr    791 embryo 08-10hr    393 embryo 10-12hr    493 embryo 12-14hr    450 embryo 14-16hr    553 embryo 16-18hr    282 embryo 18-20hr    111 embryo 20-22hr    65 embryo 22-24hr    52 larva L1    104 larva L2    80 larva L3 12hr old    36 larva L3 puffstage 1-2    76 larva L3 puffstage 3-6    446 larva L3 puffstage 7-9    905 white prepupae new    755 white prepupae 12hr    233 white prepupae 24hr    261 pupae 2d postWPP    956 pupae 3d postWPP    1191 pupae 4d postWPP    378 adult male 01day    362 adult male 05day    491 adult male 30day    487 adult female 01day    280 adult female 05day    111 adult female 30day    118 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0000241 expression level Developmental Stage   Expression Level embryo 00-02hr    39 embryo 02-04hr    218 embryo 04-06hr    312 embryo 06-08hr    791 embryo 08-10hr    393 embryo 10-12hr    493 embryo 12-14hr    450 embryo 14-16hr    553 embryo 16-18hr    282 embryo 18-20hr    111 embryo 20-22hr    65 embryo 22-24hr    52 larva L1    104 larva L2    80 larva L3 12hr old    36 larva L3 puffstage 1-2    76 larva L3 puffstage 3-6    446 larva L3 puffstage 7-9    905 white prepupae new    755 white prepupae 12hr    233 white prepupae 24hr    261 pupae 2d postWPP    956 pupae 3d postWPP    1191 pupae 4d postWPP    378 adult male 01day    362 adult male 05day    491 adult male 30day    487 adult female 01day    280 adult female 05day    111 adult female 30day    118 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    39 embryo 02-04hr  (218) embryo 04-06hr  (312) embryo 06-08hr  (791) embryo 08-10hr  (393) embryo 10-12hr  (493) embryo 12-14hr  (450) embryo 14-16hr  (553) embryo 16-18hr  (282) embryo 18-20hr  111 embryo 20-22hr    65 embryo 22-24hr    52 larva L1  104 larva L2    80 larva L3 12hr old    36 larva L3 puffstage 1-2    76 larva L3 puffstage 3-6  (446) larva L3 puffstage 7-9  (905) white prepupae new  (755) white prepupae 12hr  (233) white prepupae 24hr  (261) pupae 2d postWPP  (956) pupae 3d postWPP  (1191) pupae 4d postWPP  (378) adult male 01day  (362) adult male 05day  (491) adult male 30day  (487) adult female 01day  (280) adult female 05day  111 adult female 30day  118 Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    39 embryo 02-04hr    218 embryo 04-06hr    312 embryo 06-08hr    791 embryo 08-10hr    393 embryo 10-12hr    493 embryo 12-14hr    450 embryo 14-16hr    553 embryo 16-18hr    282 embryo 18-20hr    111 embryo 20-22hr    65 embryo 22-24hr    52 larva L1    104 larva L2    80 larva L3 12hr old    36 larva L3 puffstage 1-2    76 larva L3 puffstage 3-6    446 larva L3 puffstage 7-9    905 white prepupae new    755 white prepupae 12hr    233 white prepupae 24hr    261 pupae 2d postWPP    956 pupae 3d postWPP    1191 pupae 4d postWPP    378 adult male 01day    362 adult male 05day    491 adult male 30day    487 adult female 01day    280 adult female 05day    111 adult female 30day    118 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    39 embryo 02-04hr    218 embryo 04-06hr    312 embryo 06-08hr    791 embryo 08-10hr    393 embryo 10-12hr    493 embryo 12-14hr    450 embryo 14-16hr    553 embryo 16-18hr    282 embryo 18-20hr    111 embryo 20-22hr    65 embryo 22-24hr    52 larva L1    104 larva L2    80 larva L3 12hr old    36 larva L3 puffstage 1-2    76 larva L3 puffstage 3-6    446 larva L3 puffstage 7-9    905 white prepupae new    755 white prepupae 12hr    233 white prepupae 24hr    261 pupae 2d postWPP    956 pupae 3d postWPP    1191 pupae 4d postWPP    378 adult male 01day    362 adult male 05day    491 adult male 30day    487 adult female 01day    280 adult female 05day    111 adult female 30day    118 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    39 embryo 02-04hr    218 embryo 04-06hr    312 embryo 06-08hr    791 embryo 08-10hr    393 embryo 10-12hr    493 embryo 12-14hr    450 embryo 14-16hr    553 embryo 16-18hr    282 embryo 18-20hr    111 embryo 20-22hr    65 embryo 22-24hr    52 larva L1    104 larva L2    80 larva L3 12hr old    36 larva L3 puffstage 1-2    76 larva L3 puffstage 3-6    446 larva L3 puffstage 7-9    905 white prepupae new    755 white prepupae 12hr    233 white prepupae 24hr    261 pupae 2d postWPP    956 pupae 3d postWPP    1191 pupae 4d postWPP    378 adult male 01day    362 adult male 05day    491 adult male 30day    487 adult female 01day    280 adult female 05day    111 adult female 30day    118 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 FBgn0000241    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0000241 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 Malpighian tubules. Expression at moderate levels in the following post-embryonic organs or tissues: adult eye, adult Malpighian tubules, larval fat body. [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     Larval Malpighian Tubules no informative data Larval Fat Body no informative data Larval Salivary Gland     Larval Trachea     Larval Carcass     Adult Head no informative data Adult Eye no informative data Adult Brain     Adult Thoracic-Abdominal Ganglion     Adult Crop     Adult Midgut     Adult Hindgut     Adult Malpighian Tubules no informative data Adult Fat Body no informative data Adult Salivary Gland     Adult Heart     Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data 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 • FBgn0000241 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG17632-RB;CG17632-RA FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for bw Stages(s) 11-12 Stages(s) 13-16
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal | recessive bw5 viable bw85f4 bwD bwM58 viable, with Scer\GAL4pnr-MD237 bwGD1155 Sterility Allele fertile bwD bwM58 Other Phenotypes Allele chemical resistant | dominant | male bw1 chemical resistant | male bw1 enhancer of variegation, with lawcp1 bwD eye color defective bw1 bw2 bw2b bw2c bw4 bw5 bw24 bw30 bw33f bw33g bw37g bw38j bw45a bw47j bw53l bw69 bw72 bw75 bw81 bw609 bwA1 bwA2 bwa bwAD bwau bwB1 bwB2 bwC1 bwC4 bwC5 bwC65 bwCB bwD1 bwD114 bwD bwDa bwDe2 bwDe bwDSu169 bwE1 bwE2 bwE3 bwIG281 bwle bwM58 bwM590 bwMi59 bwPrat.PC bwR3 bwR4 bwR12 bwR14 bwR15 bwR18 bwR20 bwR27 bwR32 bwR33 bwR35 bwR40 bwR45 bwR47 bwR55 bwR56 bwR57 bwR58 bwR67 bwR68 bwR73 bwR79 bwRa bwunspecified bwV21Δ116 bwV30a bwV34k eye color defective | dominant bwD bwV1 bwV2 bwV3 bwV4 bwV5 bwV6 bwV7 bwV8 bwV29l bwV30k1 bwV30k10 bwV30k12 bwV30k18 bwV32g bwV34k bwV40b bwV54a bwV54b bwV54c bwV57e bwVD bwVDe1 bwVDe2 bwVDe3 bwVDe4 bwVI eye color defective | recessive bwDa bwDe2 bwDe bwG bwIG281 bwKN1 bwKN2 bwKN3 bwKN4 bwKN5 bwKN6 bwKN7 bwN bwR25 bwS visible bw99 bw1153 bwPrat.PC bwV1 visible | dominant bwD114 bwD bwunspecified visible | recessive bw69 bw72 bw75 bw81 bw609 bwa bwAD bwau bwCB wild-type bw4 bw24 bw59 bw+21 bwptm Phenotype manifest in Allele eye bwD bwM58 hemolymph bwR47 bwR56 ocellus bw99 pigment cell bw1 bw2 bw24 bw2b bw2c bw30 bw33f bw33g bw37g bw38j bw4 bw45a bw47j bw5 bw53l bw609 bw69 bw72 bw75 bw81 bwA1 bwA2 bwAD bwB1 bwB2 bwC1 bwC4 bwC5 bwC65 bwCB bwD bwD1 bwD114 bwDSu169 bwDa bwDe bwDe2 bwE1 bwE2 bwE3 bwG bwIG281 bwKN1 bwKN2 bwKN3 bwKN4 bwKN5 bwKN6 bwKN7 bwM58 bwM590 bwMi59 bwN bwPrat.PC bwR12 bwR14 bwR15 bwR18 bwR20 bwR25 bwR27 bwR3 bwR32 bwR33 bwR35 bwR4 bwR40 bwR45 bwR47 bwR55 bwR56 bwR57 bwR58 bwR67 bwR68 bwR73 bwR79 bwRa bwS bwV1 bwV2 bwV21Δ116 bwV29l bwV3 bwV30a bwV30k1 bwV30k10 bwV30k12 bwV30k18 bwV32g bwV34k bwV4 bwV40b bwV5 bwV54a bwV54b bwV54c bwV57e bwV6 bwV7 bwV8 bwVD bwVDe1 bwVDe2 bwVDe3 bwVDe4 bwVI bwa bwau bwle bwunspecified pigment granule bw1 wing bwD
Classical Alleles ( 138 )
For All Classical Alleles Show
Allele of bw Class Mutagen Stocks Known lesion bw1 spontaneous 2233 Yes bwD amorphic allele - genetic evidence spontaneous 65 Yes bwV1 X ray 33 Yes bwk1 14 -- bwV32g X ray 7 -- bw2b X ray 5 -- bw45a spontaneous 5 -- bw5 spontaneous 3 Yes bwVDe2 X ray 3 -- bwV34k X ray 2 -- bw16 ethyl methanesulfonate 1 Yes bw19 ethyl methanesulfonate 1 Yes bw4 spontaneous 1 -- bw75 X ray 1 -- bwv76j61 1 -- bwVDe1 X ray 1 -- bw+21 isoallele X ray 0 -- bw+ 0 -- bw108 natural population 0 -- bw111 natural population 0 -- bw1153 X ray 0 -- bw24 X ray 0 -- bw2 spontaneous 0 -- bw2c spontaneous 0 -- bw2MN ethyl methanesulfonate 0 -- bw30 spontaneous 0 -- bw33f heat-treatment 0 -- bw33g heat-treatment 0 -- bw37g spontaneous 0 -- bw38j 0 -- bw47j spontaneous natural population 0 -- bw48-366l natural population 0 -- bw49-1050 natural population 0 -- bw50b spontaneous 0 -- bw51i22 natural population 0 -- bw53l spontaneous 0 -- bw55k spontaneous 0 -- bw59 X ray 0 -- bw609 0 -- bw69 X ray 0 -- bw6 spontaneous 0 Yes bw70 X ray 0 -- bw72 X ray 0 -- bw77 X ray 0 -- bw81 X ray 0 -- bw85f4 X ray 0 Yes bw979 spontaneous 0 -- bw99 0 -- bwa spontaneous 0 -- bwAD spontaneous 0 -- bwAR ethyl methanesulfonate 0 -- bwau spontaneous 0 -- bwB1.Y spontaneous 0 Yes bwB2.Y spontaneous 0 Yes bwB3 spontaneous 0 Yes bwB4 spontaneous 0 Yes bwB5 spontaneous 0 Yes bwC65 natural population 0 -- bwCB 2-chloroethyl methanesulfonate 0 -- bwD114 X ray 0 -- bwD2 X ray 0 -- bwD3 X ray 0 -- bwDa natural population 0 Yes bwDe2 natural population 0 Yes bwDe natural population 0 Yes bwDrv12 gamma ray 0 -- bwDrv18 gamma ray 0 -- bwDSu169 ethyl methanesulfonate 0 -- bwG natural population 0 Yes bwIG281 natural population 0 Yes bwKN1 natural population 0 Yes bwKN2 natural population 0 Yes bwKN3 natural population 0 Yes bwKN4 natural population 0 Yes bwKN5 natural population 0 Yes bwKN6 natural population 0 Yes bwKN7 natural population 0 Yes bwle 0 -- bwM58 spontaneous 0 -- bwM590 0 -- bwMi59 spontaneous 0 -- bwN natural population 0 Yes bwptm natural population 0 -- bwR12 X ray 0 -- bwR14 X ray 0 -- bwR15 X ray 0 -- bwR18 X ray 0 -- bwR20 X ray 0 -- bwR25 X ray 0 -- bwR27 X ray 0 -- bwR32 X ray 0 -- bwR33 X ray 0 -- bwR35 X ray 0 -- bwR3 X ray 0 -- bwR40 X ray 0 -- bwR45 X ray 0 -- bwR47 X ray 0 -- bwR4 X ray 0 -- bwR50 X ray 0 -- bwR55 X ray 0 -- bwR56 X ray 0 -- bwR57 X ray 0 -- bwR58 X ray 0 -- bwR67 X ray 0 -- bwR68 X ray 0 -- bwR73 X ray 0 -- bwR79 X ray 0 -- bwRa 0 -- bwS natural population 0 Yes bwT50 natural population 0 Yes bwunspecified 0 -- bwV29l 226Ra alpha ray 0 -- bwV2 X ray 0 -- bwV30a X ray 0 -- bwV30k10 X ray 0 -- bwV30k12 X ray 0 -- bwV30k18 X ray 0 -- bwV30k1 X ray 0 -- bwV3 X ray 0 -- bwV40b X ray 0 -- bwV4 X ray 0 -- bwV54a gamma ray 0 -- bwV54b gamma ray neutrons 0 -- bwV54c neutrons 0 -- bwV57e X ray 0 -- bwV5 X ray 0 -- bwV6 X ray 0 -- bwV77k10(6) 0 -- bwV7 X ray 0 -- bwV8 X ray 0 -- bwVA 0 -- bwVD X ray 0 -- bwVDe3 X ray 0 -- bwVDe4 X ray 0 -- bwVI spontaneous X ray 0 -- bwW1 spontaneous 0 Yes bwW2 spontaneous 0 Yes bwx 0 --
Alleles Carried on Transgenic Constructs ( 18 )
For All Alleles Carried on Transgenic Constructs Show
Allele of bw Class Mutagen Stocks Known lesion bwGD1155 in vitro construct - RNAi in vitro construct - regulatory fusion 2 Yes bwKK103905 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes bw+t3.8 in vitro construct - genomic fragment 0 Yes bw+t30 in vitro construct - genomic fragment 0 -- bw+t8.4 in vitro construct - genomic fragment 0 -- bwA1 hypomorphic allele - genetic evidence Delta2-3 0 Yes bwA2 hypomorphic allele - genetic evidence Delta2-3 0 Yes bwB1 hypomorphic allele - genetic evidence Delta2-3 0 Yes bwB2 hypomorphic allele - genetic evidence Delta2-3 0 Yes bwC1 hypomorphic allele - genetic evidence Delta2-3 0 Yes bwC4 hypomorphic allele - genetic evidence Delta2-3 0 Yes bwC5 hypomorphic allele - genetic evidence Delta2-3 0 Yes bwD1 hypomorphic allele - genetic evidence Delta2-3 0 Yes bwE1 hypomorphic allele - genetic evidence gene conversion 0 Yes bwE2 hypomorphic allele - genetic evidence gene conversion 0 Yes bwE3 hypomorphic allele - genetic evidence gene conversion 0 Yes bwPrat.PC in vitro construct - regulatory fusion 0 Yes bwV21Δ116 hypomorphic allele - genetic evidence Delta2-3 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2R)bw5 (Dreesen et al., 1988) Df(2R)bw-DRp (Moore, 1997) Df(2R)bw-DX7 (Sass and Henikoff, 1999) Df(2R)bw-K (Koliantz, 1971) Df(2R)bw-R40 (Dreesen et al., 1988) Df(2R)bw-S46 (Moore, 1997) Df(2R)bw-W3a (Watanabe et al., 2009) Df(2R)bw-W3b (Watanabe et al., 2009) Df(2R)bw-WI320 (Moore, 1997) Df(2R)bw-WI366 (Bickel et al., 1996, Moore, 1997) Df(2R)bw-WI379 (Moore, 1997) Df(2R)bw-WI3118 (Moore, 1997) Df(2R)bw-WI3121 (Moore, 1997) Df(2R)bw-WI3128 (Moore, 1997) Df(2R)bw-WI3129 (Moore, 1997) Df(2R)bw-WI3133 (Moore, 1997) Df(2R)bw-WI3140 (Moore, 1997) Df(2R)Exel7179 (Ryder, 2004.4.26) Df(2R)or-BR6 (Sass and Henikoff, 1999) Dp(?;2)bwD (Henikoff et al., 1995, Belyaeva et al., 1997) In(2)bwD2 (Carpenter, 1994) In(2)bwR35 (Slatis, 1955) In(2LR)bw85f4 (Dreesen et al., 1988) In(2R)bwD3 (Carpenter, 1994) In(2R)bwVDe1LCyR   In(2R)CyLbwVDe1R   In(2R)CyLbwVDe2R
Duplicated in	Dp(2;2)bw-DX7 (Sass and Henikoff, 1999) Dp(2;2)Su(Pm) (Henikoff and Dreesen, 1989) Dp(2;3)P   In(2R)bwVDe2LCyR   Ts(Y;2Rt)SD-L (Lyttle, 1981)
Not disrupted in	Df(2R)3-70 (Bickel et al., 1996)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD1155} NA P{KK103905} NA characterization construct P{bw+3.8} NA P{bw+1111} NA P{bw+L.E2} NA P{bw+L.E2E1} NA P{bw+L.E3S1DE1} NA P{bw+L.E3S1E2} NA P{bw+L} NA P{bw+LR} NA P{bw+R} NA P{bwA1} NA P{bwA2} NA P{bwB1} NA P{bwB2} NA P{bwC1} NA P{bwC4} NA P{bwC5} NA P{bwD1} NA P{bwE1} NA P{bwE2} NA P{bwE3} NA P{bwV21Δ116} NA P{Prat:bw} NA
Insertions
	Type of insertions Name Expression data
Gene Ontology: Function, Process & Cellular Component ( 10 unique terms )
Terms Based on Experimental Evidence ( 1 term )
Molecular Function ( 0 terms)
Biological Process
CV term Evidence References
ommochrome biosynthetic process inferred from mutant phenotype (Tearle, 1991)
Cellular Component ( 0 terms)
Terms Based on Predictions or Assertions ( 9 terms )
Molecular Function
CV term Evidence References
ATPase activity, coupled to transmembrane movement of substances inferred from sequence or structural similarity with EMBL:AF422804 (FlyBase, 1992-) ATP binding inferred from electronic annotation with InterPro:IPR003439, InterPro:IPR017871 (FlyBase Curators et al., 2004-) eye pigment precursor transporter activity non-traceable author statement (FlyBase, 1992-, Swiss-Prot Project Members, 1989.10.1)
Biological Process
CV term Evidence References
eye pigment biosynthetic process non-traceable author statement (FlyBase, 1992-) eye pigment precursor transport traceable author statement (Reaume et al., 1991) pteridine biosynthetic process non-traceable author statement (FlyBase, 1992-) transport inferred from electronic annotation with InterPro:IPR005284 (FlyBase Curators et al., 2004-)
Cellular Component
CV term Evidence References
ATP-binding cassette (ABC) transporter complex inferred from sequence or structural similarity with EMBL:AF422804 (FlyBase, 1992-) integral to plasma membrane non-traceable author statement (Swiss-Prot Project Members, 1989.10.1)
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 bw allele Gene References bw6 w (Ewart et al., 1994, Farmer, 1974) bw38j v (Ives, 1968) bw+t8.4 Mod(Prat:bw)1-1 (Malmanche and Clark, 2003) Mod(Prat:bw)1-3 (Malmanche and Clark, 2003) Mod(Prat:bw)1-4 (Malmanche and Clark, 2003) Mod(Prat:bw)1-5 (Malmanche and Clark, 2003) Mod(Prat:bw)2-1 (Malmanche and Clark, 2003) Mod(Prat:bw)2-5 (Malmanche and Clark, 2003) Mod(Prat:bw)3-3 (Malmanche and Clark, 2003) Mod(Prat:bw)3-4 (Malmanche and Clark, 2003) Mod(Prat:bw)3-7 (Malmanche and Clark, 2003) Mod(Prat:bw)3-11 (Malmanche and Clark, 2003) Mod(Prat:bw)Y-1 (Malmanche and Clark, 2003) bwD Mod(Prat:bw)1-1 (Malmanche and Clark, 2003) Mod(Prat:bw)1-2 (Malmanche and Clark, 2003) Mod(Prat:bw)1-3 (Malmanche and Clark, 2003) Mod(Prat:bw)1-4 (Malmanche and Clark, 2003) Mod(Prat:bw)1-5 (Malmanche and Clark, 2003) Mod(Prat:bw)2-1 (Malmanche and Clark, 2003) Mod(Prat:bw)2-3 (Malmanche and Clark, 2003) Mod(Prat:bw)2-4 (Malmanche and Clark, 2003) Mod(Prat:bw)2-5 (Malmanche and Clark, 2003) Mod(Prat:bw)3-1 (Malmanche and Clark, 2003) Mod(Prat:bw)3-2 (Malmanche and Clark, 2003) Mod(Prat:bw)3-3 (Malmanche and Clark, 2003) Mod(Prat:bw)3-4 (Malmanche and Clark, 2003) Mod(Prat:bw)3-5 (Malmanche and Clark, 2003) Mod(Prat:bw)3-6 (Malmanche and Clark, 2003) Mod(Prat:bw)3-7 (Malmanche and Clark, 2003) Mod(Prat:bw)3-8 (Malmanche and Clark, 2003) Mod(Prat:bw)3-9 (Malmanche and Clark, 2003) Mod(Prat:bw)3-10 (Malmanche and Clark, 2003) Mod(Prat:bw)3-11 (Malmanche and Clark, 2003) Mod(Prat:bw)3-12 (Malmanche and Clark, 2003) Mod(Prat:bw)Y-1 (Malmanche and Clark, 2003) bwPrat.PC Mod(Prat:bw)1-1 (Malmanche and Clark, 2003) Mod(Prat:bw)1-2 (Malmanche and Clark, 2003) Mod(Prat:bw)1-3 (Malmanche and Clark, 2003) Mod(Prat:bw)1-4 (Malmanche and Clark, 2003) Mod(Prat:bw)1-5 (Malmanche and Clark, 2003) Mod(Prat:bw)2-1 (Malmanche and Clark, 2003) Mod(Prat:bw)2-3 (Malmanche and Clark, 2003) Mod(Prat:bw)2-4 (Malmanche and Clark, 2003) Mod(Prat:bw)2-5 (Malmanche and Clark, 2003) Mod(Prat:bw)3-1 (Malmanche and Clark, 2003) Mod(Prat:bw)3-2 (Malmanche and Clark, 2003) Mod(Prat:bw)3-3 (Malmanche and Clark, 2003) Mod(Prat:bw)3-4 (Malmanche and Clark, 2003) Mod(Prat:bw)3-5 (Malmanche and Clark, 2003) Mod(Prat:bw)3-6 (Malmanche and Clark, 2003) Mod(Prat:bw)3-7 (Malmanche and Clark, 2003) Mod(Prat:bw)3-8 (Malmanche and Clark, 2003) Mod(Prat:bw)3-9 (Malmanche and Clark, 2003) Mod(Prat:bw)3-10 (Malmanche and Clark, 2003) Mod(Prat:bw)3-11 (Malmanche and Clark, 2003) Mod(Prat:bw)3-12 (Malmanche and Clark, 2003) Mod(Prat:bw)Y-1 (Malmanche and Clark, 2003) bwT50 w (Ewart et al., 1994) bwx ade2 (Tiong et al., 1989) unspecified st (Campbell and Nash, 2001)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 63324 DroID - A comprehensive database of gene and protein interactions. • FBgn0000241 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 37724
Orthologs
Genome-wide drosophilid orthologs	Dana\GF11841 Dere\GG20023 Dgri\GH20650 Dmoj\GI19015 Dper\GL17554 Dpse\GA24827 Dsec\GM15535 Dsim\GD25039 Dvir\bw Dwil\GK23049 Dyak\GE11560
Curated drosophilid orthologs	Dana\Pm (Sturtevant and Novitski, 1941) Dana\pe   Dbuz\bw (Pla et al., 1986) Dmas\pp (Price, 1954) Dmau\bw   Dpro\Pm (Spassky et al., 1950) Dpse\GA14586   Dpse\pr (Sturtevant and Tan, 1937) Dsim\bw (Rux and Coyne, 1991) Dvir\bw   Dvir\es (Sturtevant and Novitski, 1941, Price, 1954) Dwil\bw (Spassky and Dobzhansky, 1950)
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AGAP007655-PA Anopheles gambiae (mosquito) XP_395665.3 Apis mellifera (honey bee) CPIJ012805 Culex pipens (mosquito) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • FBgn0229360 FBgn0225011 FBgn0013916 FBgn0196351 FBgn0170453 FBgn0246212 FBgn0155157 FBgn0141754 FBgn0128113 FBgn0112222 FBgn0088881 CPIJ012805 AGAP007655 AAEL017188
Stocks & Reagents
Stocks Listed in FlyBase ( 2369 )
Bloomington	245 bw1 248 bw2b 9474 bw19 9475 bw16 264 cn1 bw1 433 vg1 bw1 668 v1; bw1
Kyoto	101339 bwD 103250 bw1 105812 bw1 101540 lt1 bw1
VDRC	v101584 P{KK103905}VIE-260B
	More stocks available...
Genomic Clones ( 2 )
	BACR25I01 BACR48M01 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 140 )
	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	GH12107 GM07634 IP02562 LD46606
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	GH09183 GM03224 GM03815 GM05045 LP10226 RE14280 RE28425 RE35814 RE35826 RE45468 RE50035 RE51837 RH10564 SD04181 SD08607 SD10309 SD10954 SD21404
Other clones	12904 16872 33807 73497 120554 167314 214383.66 CK01.93C.D2 CK00276 CK00290 CK01013 CK01418 CK01435 CK01710 CK02129 CK02242 CK02431 EK001146 EK001816 EK048241 EK050126 EK050902 EK051431 EK058816 EK067046 EK072945 EK074010 EK080748 EK085844 EK085928 EK086522 EK087521 EK088646 EK090460 EK098818 EK104826 EK109812 EK110627 EK114255 EK114502 EK119406 EK121537 EK122408 EK128633 EK130555 EK132303 EK136345 EK145861 EK146701 EK150642 EK154733 EK157335 EK164553 EK165633 EK166523 EK168134 EK169053 EK184348 EK184436 EK190542 EK191430 EK199217 EK201143 EK213058 EK220032 EK226934 EK235206 EK242629 EK244018 EK247539 EK249423 EK254438 EK261620 EK261662 EK265103 EK266014 EK266032 EK269517 EK272035 EK272411 EK272734 EK279364 EK283053 EK288824 EK298018 EK299637 EK302060 EK302235 EK306316 EK307114 EN12547 EN16847 EP07313 EP09126 EP09949 EP10433 IP05438 RP004040543 RP004071103 RP004126961 RP004164906 RP004179572 RP004211026 RP004230664 RP004235487 RP004237933 RP004244546 RP004247884 RP004251136 RP004416667 RP004451739 RP004468523 RP004469280 RP004483308 RP004533018 RP004545347 RP004556901 RP004557330
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0000241 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 37724
Antibody Information
Other Information
Discoverer
	Waaler, 15th Oct. 1919.
Etymology
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of
Additional comments	Alleles designated bwR: brown-Rearranged were X-ray-induced derivatives of bw or bw+ by Slatis, 48k16. (Slatis, 1955). They exhibit variegated phenotype in combination with +. Mostly homozygous lethal; survivors have brown eyes. More complete descriptions in Lindsley and Grell, 1968. All associated with chromosome rearrangements with one breakpoint in 59DE and one in proximal heterochromatin.
Other Comments
	bw can be simultaneously acted upon by pericentric and distal blocks of heterochromatin. (Sass and Henikoff, 1999) An element, P{Prat:bw} encoding a very low pigment level has been used to generate tandem arrays of up to 19 transposon copies at cytological location 65A10. The tandem arrays have altered polytene chromosome morphology. (Clark et al., 1998) One of a class of genes with TATA-less promoters that have a subset of the conserved DPE sequence. (Burke and Kadonaga, 1996) Fluorescent in situ hybridisation (FISH) to interphase diploid nuclei demonstrates that the insertion of heterochromatin into bw causes the aberrant association of bw and its homologous copy with heterochromatin. The cytological association of the heterochromatic region is affected by chromosomal distance from heterochromatin and by genic modifiers of PEV. (Csink and Henikoff, 1996) Studies of PEV using a transposon-borne bw allele support a model in which local alterations affect the affinity of a gene region for nearby heterochromatin via homology-based pairing, rather than heterochromatin being propagated along the chromosome. (Sabl and Henikoff, 1996) Transcripts of bw, st and w are all modulated by Inr-b in a developmentally-specific manner. (Bhadra et al., 1995) Dp(?;2)bwD position effect variegation (PEV) is observed as the very strong dominant variegating effect on the bw+ copy present on the homologous chromosome. Dp(?;2)bwD fails to coalesce with the chromocenter when its position along the chromosome places it beyond a threshold distance from heterochromatin. This threshold depends upon configuration of Dp(?;2)bwD chromosome and its paired homolog. The basis of silencing is paractivation across paired copies of the duplication as opposed to cis-spreading along the chromosome. (Henikoff et al., 1995) The D.virilis and D.melanogaster bw genes show functional conservation: Dvir\bw+ rescues the D.melanogaster mutant bw phenotype. Sequences required for transactivation are also conserved as Dvir\bw in D.melanogaster shows dominant PEV. (Martin-Morris and Henikoff, 1995) Dvir\bw transformed into D.melanogaster can be expressed. Induction of a PEV mutation in the Dvir\bw transgene reveals that Dvir\bw is capable of being trans-inactivated in D.melanogaster. These results support the conclusion that sequences responsible for trans-inactivation are conserved to regulate bw expression. (Martin-Morris and Henikoff, 1995) A fifth of mutations selected on the basis of suppression of bwD carry rearrangements that modify PEV by changing the chromosomal position of bwD, suggesting that PEV depends on nuclear position. (Talbert et al., 1994) The sequence-specific component of the bw locus that is responsible for trans-inactivation may be the site at which a bw-specific transcription factor binds. Deletion analysis of the 8.4kb EcoRI fragment carrying the wild type bw+t8.4 from the P{bw+L}92C construct demonstrates that a 3.8kb fragment containing the bw coding region and 850bp of upstream sequences includes the genetic element responsible for trans-inactivation. (Martin-Morris et al., 1993) Chromatographic pattern of pteridine eye pigments neodrosopterin, sepiapterin, pterin, aurodrosopterin, acetyldihidrohomopterin, isoxanthopterin, biopterin and drosopterin measured in bw alleles. (Millan and Najera, 1992) Analysis of variance of developmental time and viability of pteridine pathway mutants in sf, se, Hn, dke and bw, indicated that viability of induced and natural population alleles is the same whereas developmental time tends to be longer for induced mutations as compared to natural population alleles. (Millan and Najera, 1992) Trans inactivation at bw requires some feature or component of heterochromatin, somatic pairing between variegating and wild-type alleles and a sequence in or near the bw gene in trans to the source of the heterochromatin. 35 lines carrying ectopic copies of bw were examined in the presence of dominant "bwV" alleles: no mutant spots or clones characteristic of trans-inactivation were seen. One transformant, "R26", has the P element construct colocalized with the resident bw+ and was trans-inactivated, suggesting that trans-inactivation is a chromosome-local phenomenon. Induction of position effect variegation on an ectopic copy of bw by chromosome rearrangement causes trans-inactivation of its paired homolog. Excisions and deletions of the copy of bw in cis to the rearrangement partially suppress transinactivation in a size-dependent manner, as if local somatic pairing (disrupted by the deletions) is necessary for trans-inactivation to occur. (Dreesen et al., 1991) bw has no substantial affect on ommochrome pigmentation in the fat body, ocelli or tubules. (Tearle, 1991) Ultrastructure and Ca-sequestering properties of eye colour pigment granules (PGs) are studied in the eyes of wild type and mutant flies. A new type of ommochrome PG localises in primary pigment cells and basal terminals of the processes of Semper cells. At room temperature these PGs exhibit OsO4-dependent structural ability. X ray microanalysis revealed a less Ca-binding ability of the PGs as compared to those in secondary pigment cells. (Ukhanov, 1990) Transinactivation at the bw gene is antimorphic, occurs at the level of mRNA accumulation and is sensitive to certain structural configurations of the bw chromosomal region. (Henikoff and Dreesen, 1989) A bw cDNA has been cloned and sequenced. (Dreesen et al., 1988) Mutant alleles are useful as markers in clonal analysis. (Lawrence et al., 1986) Uptake studies indicate that bw, like w, blocks the transport of guanine and xanthine, likely pteridine precursors (Sullivan and Sullivan, 1975). Sequence comparisons between the predicted bw- and w-encoded proteins show that they are similar to one another and to subunits of active transport family members (Mount, 1987; Dreesen, Johnson and Henikoff, 1988).
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_079100 NM_206206 RefSeq (Proteins) • NP_523824 NP_995928 DDBJ / EMBL / GenBank • AAA28397 AAA28398 AAC37214 AAF47020 AAF47020.3 AAM50157 AAS64760.1 AAS64761 AAS64761.1 AF047460 AI134596 AW940450 AY119503 L05635 L23543 M20630 Z50313 Entrez Gene - A searchable database of RefSeq genes. • 37724 UniProtKB/Swiss-Prot • P12428
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • ABC transporter-like (IPR003439) ATPase, AAA+ type, core (IPR003593) ABC-2 type transporter (IPR013525) Pigment precursor permease (IPR005284) ABC transporter, conserved site (IPR017871)
Linkouts
	BioGRID - A database of protein and genetic interactions • 63324 DroID - A comprehensive database of gene and protein interactions. • FBgn0000241 DRSC - Results from RNAi screens. • FBgn0000241 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0000241 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG17632-RB;CG17632-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0000241 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 37724 InParanoid A subset of ortholog calls from InParanoid. • AGAP007655-PA Anopheles gambiae (mosquito) XP_395665.3 Apis mellifera (honey bee) CPIJ012805 Culex pipens (mosquito) InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 37724 modMine - Data generated by the modENCODE project. • FBgn0000241 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • FBgn0229360 FBgn0225011 FBgn0013916 FBgn0196351 FBgn0170453 FBgn0246212 FBgn0155157 FBgn0141754 FBgn0128113 FBgn0112222 FBgn0088881 CPIJ012805 AGAP007655 AAEL017188
Synonyms & Secondary IDs ( 13 )
Reported As
Symbol Synonym	bw (Sweeney et al., 2008, Watanabe et al., 2009, Nisha et al., 2008, Talbert and Henikoff, 2006, Beumer et al., 2006, Nisha and Csink, 2007, Zuckerkandl and Cavalli, 2007, Wang et al., 2004, Stewart et al., 2005, Engel, 2008, Schneiderman and Ahmad, 2008, Lindsley and Grell, 1968, Sage et al., 2005, Morrow et al., 2005, Harmon and Sedat, 2005, Thakar and Csink, 2005, Thakar et al., 2006, Lipatov et al., 2005, Anaka et al., 2008, Pasyukova et al., 2004, Meyer et al., 2008, Schneiderman et al., 2010, Enright et al., 2003) CG17632 (Dean et al., 2001, Keleman et al., 2009.8.5, ) Pm (Goldschmidt, 1999) ptm   Su(wco2)   Su(wcoJ) (Farmer, 1974) unnamed (Oliver, 1937)
Name Synonym	brow (Schneiderman et al., 2010) brown (Platero, 1998.2.5, Nitasaka, 1992, Kibanov et al., 2008, Schneiderman and Ahmad, 2008, Sage et al., 2005, Morrow et al., 2005, Harmon and Sedat, 2005, Thakar and Csink, 2005, Lipatov et al., 2005, Anaka et al., 2008, Borycz et al., 2008, Pasyukova et al., 2004, Enright et al., 2003) Plum (Greenspan, 2003, Morgan, 1947) pteridine-modifier   Suppressor of white-coral-2   Suppressor of white-coral J (Farmer, 1974)
Secondary FlyBase IDs
References ( 308 )
Generate a list of	All references relating to this gene ( 308 )
List References by type	Research paper  ( 191 ) Supplementary material  ( 2 ) Review  ( 54 ) Personal communication to FlyBase  ( 9 ) Abstract  ( 29 ) FlyBase analysis  ( 2 ) DNA/RNA sequence record  ( 2 ) Teaching note  ( 1 ) Book  ( 4 ) Protein sequence record  ( 1 ) Stock list  ( 4 ) Computer file  ( 1 ) Thesis  ( 2 ) Interview  ( 1 ) Letter  ( 1 ) Automatic genome annotation  ( 1 ) Curated genome annotation  ( 1 ) Conference report  ( 1 )
Recent research papers ( 2 )
	Ambegaokar and Jackson, 2010, Genetics 186(1): 435--442 Interaction Between Eye Pigment Genes and Tau-Induced Neurodegeneration in Drosophila melanogaster. [FBrf0211818] Schneiderman et al., 2010, PLoS Genet. 6(9): e1001095 Perturbation Analysis of Heterochromatin-Mediated Gene Silencing and Somatic Inheritance. [FBrf0211786] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...