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Gene Dmel\en

General Information
Symbol	Dmel\en	Species	D. melanogaster
Name	engrailed	Annotation symbol	CG9015
Feature type	protein_coding_gene	FlyBase ID	FBgn0000577
Gene Model Status	Current	Stock availability	38 publicly available
Also Known As	Eng, en1
Genomic Location
Chromosome (arm)	2R	Recombination map	2-62.0
Cytogenetic map	47F17-48A1	Sequence location	2R:7,411,509..7,415,715 [-]
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 engrailed is referred to in FlyBase by the symbol Dmel\en (CG9015, FBgn0000577). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in negative regulation of transcription; RNA polymerase II distal enhancer sequence-specific DNA binding. There is experimental evidence that it is involved in the biological process: anterior commissure morphogenesis; neuroblast fate determination; axon guidance; spiracle morphogenesis, open tracheal system; gonad development; negative regulation of gene expression; ventral midline development; negative regulation of transcription from RNA polymerase II promoter; genital disc development; genital disc anterior/posterior pattern formation. 184 alleles are reported. The phenotypes of these alleles are annotated with: organ system subdivision; adult segment; abdominal ventral denticle belt; adult mesothoracic segment; multi-cell-component structure; organ system; nervous system; presumptive embryonic/larval nervous system; embryonic/larval neuron; antennal segment. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: Helix-turn-helix motif, lambda-like repressor; Homeobox; Homeobox engrailed; Homeobox engrailed, C-terminal; Homeobox engrailed-type, conserved site; Homeobox, conserved site; Homeobox, eukaryotic; Homeodomain-like; Homeodomain-related. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of very low expression. Peak expression observed within 00-12 hour embryonic stages. Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: larval hindgut. Comments on Affy2 ProbeSet: ProbeSet 1627445_s_at completely aligns to an exonic region common to each of the 2 FlyBase-annotated transcript isoforms of en. Gene sequence location is 2R:7411509..7415715.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	en: engrailed (T.Kornberg) en: engrailed From Eker, 1929, Hereditas 12: 217-22. Four classes of alleles, all recessive. (1) en1. Viable hemizygous and homozygous; fertile. Longitudinal cleft extends from rear border of scutellum forward, may be reduced to median nick or posterior flattening of scutellum. Wings larger, broader, and thin textured with spatulate end; venation and distribution of sensilla abnormal in posterior wing compartment. Variable duplication of anterior triple row bristles on posterior margin; alula reduced, with costal-like bristles. In males, extra sex comb often present (Brasted, 1941, Genetics 26: 347-73), smaller than normal, and in mirror-image position in posterior compartment. Duplications of transverse rows in female prothoracic leg, extra bristles in mesothoracic and metathoracic tarsi (Garcia-Bellido, and Santamaria, 1972, Genetics 72: 87-104; Lawrence, Struhl, and Morata, 1979, J. Embryol. Exp. Morph. 51: 195-208). Action of en1 is autonomous except for scutellar cleft (Tokunaga, 1961, Genetics 46: 157-76; Stern and Tokunaga, 1968, Proc. Nat. Acad. Sci. USA 60: 1252-59; Garcia-Bellido, and Santamaria, 1972, Genetics 72: 87-104). Clones of en cells of posterior compartment origin fail to respect anterior-posterior compartment border in wing disc as do mwh clones in wing discs of en1 homozygotes (Morata and Lawrence, 1975, Nature 255: 614-17; Morata and Lawrence, 1976, Dev. Biol. 50: 321-37). en abnormalities are associated with posterior compartment structures only, except for scutellar cleft. Restriction of glucose-6-phosphate dehydrogenase, 6-phosphogluconate dehydrogenase (Cunninghamn, Smith, Makowski, and Kuhn, 1983, Mol. Gen. Genet. 191: 238-43) and of a protein recognized by monoclonal antibody PS2 (Brower, 1984, Nature 310: 496-98; 1987, EMBO J. 5: 2649-56) to posterior compartment of wing disc altered by en1. Interaction with ci, cg, sx (House, 1953, Genetics 38: 200-15; House, 1953, Genetics 38: 309-27; House, 1961, Genetics 46: 871; Mukherjee, 1965, Genetics 51: 285-304; Datta and Mukherjee, 1971, Genetics 68: 269-86) and fu (Fausto-Sterling and Smith-Schiess, 1982, EMBO J. 1: 827-33) partially increase phenotype. No suppression by su(Hw). (2) Lethal alleles with normal cytology. Embryonic lethal. Anterior margin of each segment defective. Pair rule defects in naked cuticle of T1, T3, A2, A4, A6, A8 result in pair-wise fusion of adjacent segments. Autonomous effects in adult cuticular clones observed in posterior compartment of proboscis, thorax, abdomen, and genitalia. enlethal clones are without effect in anterior compartments and in eye-antennal region (Kornberg, 1981, Proc. Nat. Acad. Sci. USA 78: 1095-99; Kornberg, 1981, Dev. Biol. 86: 363-72; Lawrence and Struhl, 1982, EMBO J. 1: 827-33). The en1/enlethal heterozygote characterized by wing abnormalities only; disruption of anterior crossvein, gap in vein IV, and occasional socketted bristles on posterior margin. In some combinations complementation is complete or nearly so (Condie and Brower, 1989, Dev. Biol. 135: 31-42). No maternal effect (Lawrence, Johnston and Struhl, 1983, Cell 35: 27-34). (3) Deficiencies and lethal alleles with inversion or translocation breakpoints. Embryonic lethal. Embryonic segment defects slight and variable. Alleles of this class in heterozygous combination witn en1 produce adults more extreme than en1. For example, in en1/en2, legs are truncated, the tarsi reduced to densely bristled stumps; wings are greatly enlarged and spatulate with greater disruption of veins IV and V; higher penetrance of socketed bristles along the posterior margin. Extreme scutellar cleft. At 29, duplication of anterior compartment bristles in mirror-image symmetry in posterior compartment of second antennal segment (Morata, Kornberg, and Lawrence, 1983, Dev. Biol. 99: 27-33). (4) Non-lethal alleles with breakpoints. Hemizygous viable, embryos normal. Heterozygous with other allele classes, variable gaps in wing veins IV and V. Variable reductions or deletions in male and female genitalia (Epper and Sanchez, 1983, Dev. Biol. 100: 387-98). Scutellum may also be affected. spt: spermatheca At 28, female has two spermathecae but ducts partly fused; at 25, only one enlarged spermatheca on one duct; at 18, a duct with three branches, each bearing a spermatheca. Temperature-sensitive period in third larval instar. Female fertility not greatly affected. RK3.
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 BKN45260 HFA07609 BKN27874 References McKnight et al., 2011, Mol. Cell. Biol. 31(23): 4746--4759 Kazemian et al., 2010, PLoS Biol. 8(8): Tchuraev and Galimzyanov, 2009, J. Theor. Biol. 259(4): 659--669 Rand et al., 2010, Insect Biochem. Mol. Biol. 40(11): 792--804 Slack et al., 2006, BMC Genet. 7: 33 Controlled Vocabulary Terms negative regulation of transcription from RNA polymerase II promoter ventro-lateral serotonergic neuron negative regulation of gene expression RNA polymerase II distal enhancer sequence-specific DNA binding RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in negative regulation of transcription
FB2011_10	References Wang et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(27): 11139--11144 Gurunathan et al., 2004, BMC Bioinformatics 5: 202 Grad et al., 2004, Bioinformatics 20(16): 2738--2750 González et al., 2008, Bioinformatics 24(16): i234--i240 Jia et al., 2010, J. Biol. Chem. 285(48): 37218--37226 Babcock et al., 2011, Curr. Biol. 21(18): 1525--1533 Wojcinski et al., 2011, Dev. Biol. 358(1): 168--180
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 47F17-48A1   Limits computationally determined from genome sequence between P{lacW}Tapδk17005&P{lacW}k05103 and P{lacW}l(2)k13403k13403&P{lacW}Egmk14708
Experimentally Determined Cytological Location
Cytogenetic map Notes References 48A-48A   (determined by in situ hybridisation)   (Brand, 1997.6.30) 48A-48A   (determined by in situ hybridisation)   (Coleman et al., 1987) 48A-48A   (determined by in situ hybridisation)   (Drees et al., 1987) 48A2-48A2     (Tearle and Nusslein-Volhard, 1987)
Experimentally Determined Recombination Data
Location	2-62.0   2-64.0 (Kornberg, 1981) 2-62 (Tearle and Nusslein-Volhard, 1987) 2- (Hadorn and Graber, 1946)
Left of (cM)	L (Hadorn and Graber, 1946) vg (Kornberg, 1981)
Right of (cM)	cn (Kornberg, 1981) lt (Hadorn and Graber, 1946)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\en for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) en-RA FBtr0088096  NM_078976   2467   552 en-RB FBtr0088095  NM_165841   2794   552
Additional Transcript Data & Comments
Reported size (kB)	3.6, 2.7, 1.4 (northern blot) (Drees et al., 1987)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein en-PA   FBpp0087198   59.4   552   7.07     NP_523700   AAF58639 en-PB   FBpp0087197   59.4   552   7.07     NP_725059   AAM68711
Additional Polypeptide Data & Comments
Reported size (kDa)
Comments	en protein is used as a marker for the embryonic dorsal large intestine. (Maeda et al., 2007) Polyclonal antibody is to the entire en protein. (Chanas et al., 2004) en expression in the epidermis, but not the central nervous system, is disrupted in embryos which over express bcd. This disruption results in either the loss or fusion of en stripes. (Namba et al., 1997) Two monoclonal antibodies were generated against en (and also recognize inv), designated 4D9 and 4F11. 4D9 is directed to an engrailed specific region of the homeodomain, and recognizes en protien in several different species, but does not cross react with other homeodomain proteins. (Patel et al., 1989)
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Homeobox, conserved site (IPR017970) Helix-turn-helix motif, lambda-like repressor (IPR000047) Homeobox engrailed (IPR000747) Homeobox (IPR001356) Homeodomain-like (IPR009057) Homeodomain-related (IPR012287) Homeobox engrailed, C-terminal (IPR019549) Homeobox engrailed-type, conserved site (IPR019737) Homeobox, eukaryotic (IPR020479) PDB - Protein Data Bank. An information portal to biological macromolecular structures • 1ENH 1HDD 2HDD 3HDD
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA441254   AE013599 AAF58639   AE013599 AAM68711   AW942397   AY069448 AAL39593   AY184070   AY184071   AY184072   AY184073   AY184074   AY184075   AY184076   AY184077   AY184078   AY184079   AY184080   AY184081   BF492050   BI627021   CO177317   CO189044   CO189206   CO263390   CO292199   CO293330   CO294005   CO301541   CO339817   CO340604   EC059297   EC064318   EC069653   EC080757   EC082573   EC085693   EC198144   EC202055   EC229837   EC235553   EC242702   EC247271   EC254515   EC255864   EL877571   GH941822   GH941931   GH941973   GH942642   GH943951   GH944634   GH945168   GH945255   GH946300   GH948457   GH949880   GH950067   GH950395   GH951149   GH952236   GH952297   GH953458   GH953993   GH954466   GH954811   GH956430   GH956645   GH957151   GH957416   GH957657   GH957934   GH958915   GH959950   GH960531   GH960548   GH961138   GH961711   GH962150   GH963189   GH964159   GH964234   GH965025   GH965510   GH966398   GH966736   GH967991   GH968091   GH969551   GH969702   GH971140   GH972677   GH973058   GH973853   GH974196   GH974646   GH975304   GH975337   GH976141   GH976157   GH976178   GH976346   GH976753   GH977197   GH978656   GH978835   GH979091   GH979129   GH979607   GH979640   GH979770   GH980353   GH981098   GH981471   GH981575   GH982385   GH982772   K03055   K03056   M10017 AAA65478   M29285   X01765 CAA25906
UniProtKB/Swiss-Prot	P02836
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 insertion site P{en-lacZ(Xho)}en[Xho25] 2R:7,415,645..7,415,645 comment=Annotated insertion location based on author report that insertion maps 257 nucleotides upstream of the en transcription start. The transcription start is from FBrf0064711. The orientation of the insertion was not reported. evidence=experimental (Gustavson et al., 1996) insertion site P{en1}en[17en52] 2R:7,413,864..7,415,368 comment=The insertion maps to the first exon of en. The orientation of the insertion was not reported. evidence=experimental (Mellerick and Nirenberg, 1995) protein binding site FBsf0000159742 2R:7,416,312..7,416,322 bound_moiety=hb-XP evidence=experimental (Zuo et al., 1991, Bergman et al., 2004) protein binding site FBsf0000160045 2R:7,416,281..7,416,316 bound_moiety=en-XP bound_moiety=eve-XP comment=Binding site k1 in the en upstream region is bound by eve and zen and weakly by prd and en. eve-binding site k1 contains three tandem repeats similar to the consensus sequence TCAATTAAAT. evidence=experimental bound_moiety=prd-XP (Bergman et al., 2004, DiNardo et al., 1988, Desplan et al., 1988, Han et al., 1989, Hoey and Levine, 1988) protein binding site FBsf0000160046 2R:7,416,263..7,416,272 bound_moiety=hb-XP evidence=experimental (Bergman et al., 2004, Zuo et al., 1991) protein binding site FBsf0000161131 2R:7,416,242..7,416,262 bound_moiety=zen-XP bound_moiety=en-XP evidence=experimental comment=Binding site k2 in the en upstream region is bound by eve and zen and weakly by prd and en. eve-binding site k2 contains an inverted repeat similar to the consensus sequence TCAATTAAAT. bound_moiety=eve-XP (DiNardo et al., 1988, Desplan et al., 1988, Hoey and Levine, 1988, Bergman et al., 2004) protein binding site FBsf0000160047 2R:7,416,246..7,416,255 bound_moiety=Kr-XP evidence=experimental (Zuo et al., 1991, Bergman et al., 2004) protein binding site FBsf0000160048 2R:7,416,118..7,416,143 comment=Binding site k3 in the en upstream region is bound by eve and zen and weakly by prd and en. evidence=experimental bound_moiety=en-XP bound_moiety=eve-XP bound_moiety=en-XP (Bergman et al., 2004, DiNardo et al., 1988, Hoey and Levine, 1988) protein binding site FBsf0000160049 2R:7,413,649..7,413,659 bound_moiety=ftz-XP comment=ftz-binding site cad1 evidence=experimental (Florence et al., 1997, Bergman et al., 2004) protein binding site FBsf0000160050 2R:7,413,637..7,413,647 bound_moiety=ftz-f1-XP comment=ftz-f1-binding site nhr1 evidence=experimental (Florence et al., 1997, Bergman et al., 2004) protein binding site FBsf0000160051 2R:7,413,625..7,413,634 bound_moiety=ftz-XP comment=ftz-binding site hd1 evidence=experimental (Bergman et al., 2004, Florence et al., 1997) protein binding site FBsf0000160052 2R:7,413,477..7,413,486 bound_moiety=ftz-XP comment=ftz-binding site cad2 evidence=experimental (Florence et al., 1997, Bergman et al., 2004) protein binding site FBsf0000160053 2R:7,413,412..7,413,422 bound_moiety=ftz-f1-XP comment=ftz-f1-binding site nhr2 evidence=experimental (Florence et al., 1997, Bergman et al., 2004) protein binding site FBsf0000160054 2R:7,413,234..7,413,243 bound_moiety=ftz-XP comment=ftz-binding site hd2 evidence=experimental (Florence et al., 1997, Bergman et al., 2004) protein binding site FBsf0000160055 2R:7,413,222..7,413,231 evidence=experimental bound_moiety=ftz-XP comment=ftz-binding site hd3 (Florence et al., 1997, Bergman et al., 2004) protein binding site FBsf0000160056 2R:7,413,211..7,413,220 bound_moiety=ftz-XP comment=ftz-binding site cad3 evidence=experimental (Florence et al., 1997, Bergman et al., 2004) protein binding site FBsf0000160057 2R:7,413,122..7,413,131 bound_moiety=ftz-XP comment=ftz-binding site hd4 evidence=experimental (Florence et al., 1997, Bergman et al., 2004) protein binding site FBsf0000160058 2R:7,413,089..7,413,098 bound_moiety=ftz-XP comment=ftz-binding site hd5 evidence=experimental (Bergman et al., 2004, Florence et al., 1997) protein binding site FBsf0000161132 2R:7,413,080..7,413,099 bound_moiety=en-XP evidence=experimental (Bergman et al., 2004, Kassis et al., 1989, Desplan et al., 1988) protein binding site FBsf0000160059 2R:7,413,020..7,413,029 bound_moiety=ftz-XP comment=ftz-binding site hd6 evidence=experimental (Bergman et al., 2004, Florence et al., 1997) protein binding site FBsf0000160060 2R:7,413,019..7,413,028 bound_moiety=en-XP evidence=experimental (Kassis et al., 1989, Desplan et al., 1988, Bergman et al., 2004) protein binding site FBsf0000160061 2R:7,412,991..7,413,000 bound_moiety=ftz-XP comment=ftz-binding site hd7 evidence=experimental (Florence et al., 1997, Bergman et al., 2004) protein binding site FBsf0000161133 2R:7,412,987..7,412,998 bound_moiety=en-XP evidence=experimental (Kassis et al., 1989, Desplan et al., 1988) protein binding site FBsf0000161017 2R:7,415,454..7,415,471 evidence=experimental (Soeller et al., 1988, Bergman et al., 2004) protein binding site FBsf0000161018 2R:7,415,423..7,415,443 evidence=experimental (Soeller et al., 1988, Bergman et al., 2004) protein binding site FBsf0000161019 2R:7,415,527..7,415,541 evidence=experimental (Soeller et al., 1988, Bergman et al., 2004) protein binding site FBsf0000161020 2R:7,415,475..7,415,489 evidence=experimental (Soeller et al., 1988, Bergman et al., 2004) protein binding site FBsf0000161021 2R:7,415,652..7,415,672 evidence=experimental (Bergman et al., 2004, Soeller et al., 1988) protein binding site FBsf0000161022 2R:7,415,693..7,415,735 evidence=experimental (Bergman et al., 2004, Soeller et al., 1988) protein binding site FBsf0000161023 2R:7,415,570..7,415,586 evidence=experimental (Soeller et al., 1988, Bergman et al., 2004) protein binding site FBsf0000161024 2R:7,415,618..7,415,638 evidence=experimental (Soeller et al., 1988, Bergman et al., 2004) protein binding site FBsf0000161025 2R:7,416,283..7,416,314 bound_moiety=ftz-XP evidence=experimental (Desplan et al., 1988, Bergman et al., 2004) protein binding site FBsf0000161026 2R:7,416,240..7,416,261 bound_moiety=ftz-XP evidence=experimental (Desplan et al., 1988, Bergman et al., 2004) protein binding site FBsf0000159966 2R:7,416,241..7,416,262 bound_moiety=en-XP comment=Binding site k2 in the en upstream region is bound by eve and zen and weakly by prd and en. eve-binding site k2 contains an inverted repeat similar to the consensus sequence TCAATTAAAT. evidence=experimental (DiNardo et al., 1988, Desplan et al., 1988) protein binding site FBsf0000159963 2R:7,413,081..7,413,098 bound_moiety=en-XP evidence=experimental (Kassis et al., 1989, Desplan et al., 1988) protein binding site FBsf0000159964 2R:7,412,986..7,412,998 bound_moiety=en-XP evidence=experimental (Kassis et al., 1989, Desplan et al., 1988) regulatory region FBsf0000161288 2R:7,415,693..7,415,735 comment=en upstream region (H) bound by factor(s) in a 0-12hr embryonic extract. evidence=experimental (Soeller et al., 1988) regulatory region FBsf0000161290 2R:7,415,651..7,415,671 comment=en upstream region (G) bound by factor(s) in a 0-12hr embryonic extract. evidence=experimental (Soeller et al., 1988) regulatory region FBsf0000161289 2R:7,415,618..7,415,638 comment=en upstream region (F) bound by factor(s) in a 0-12hr embryonic extract. evidence=experimental (Soeller et al., 1988) regulatory region FBsf0000161291 2R:7,415,570..7,415,586 comment=en upstream region (E) bound by factor(s) in a 0-12hr embryonic extract. evidence=experimental (Soeller et al., 1988) regulatory region FBsf0000161283 2R:7,415,527..7,415,540 comment=en upstream region (D) bound by factor(s) in a 0-12hr embryonic extract. evidence=experimental (Soeller et al., 1988) regulatory region FBsf0000161284 2R:7,415,476..7,415,489 comment=en upstream region (C) bound by factor(s) in a 0-12hr embryonic extract. evidence=experimental (Soeller et al., 1988) regulatory region FBsf0000161277 2R:7,415,454..7,415,471 comment=en upstream region (B) bound by factor(s) in a 0-12hr embryonic extract. evidence=experimental (Soeller et al., 1988) regulatory region FBsf0000161278 2R:7,415,423..7,415,443 comment=en upstream region (A) bound by factor(s) in a 0-12hr embryonic extract. evidence=experimental (Soeller et al., 1988) regulatory region FBsf0000161274 2R:7,415,348..7,415,389 comment=en minimal promoter evidence=experimental (Orihara et al., 1999) regulatory region FBsf0000161285 2R:7,415,350..7,415,356 comment=Downstream promoting element evidence=experimental (Orihara et al., 1999) regulatory region FBsf0000161286 2R:7,412,823..7,413,889 comment=The intron fragment confers striped expression in early embryos. Expression also occurs in the progenitor of the fat body and in some neurons. evidence=experimental (Kassis, 1990)
External Data
Linkouts
Crossreferences	EPD - Eukarytoic Promoter Database, an annotated collection of POL II promoters • 24009
Expression Data
Transcript Expression
No Assay Recorded Stage Tissue/Position (including subcellular localization) Reference larval stage   wing disc   (Couso et al., 1993) haltere disc   (Couso et al., 1993) ventral thoracic disc   (Couso et al., 1993) third instar larval stage   ventral thoracic disc | restricted   (Masucci et al., 1990) in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage   organism | segment polarity expression pattern   (Hughes and Krause, 2001) embryonic stage 5   organism | 60% egg length   (Tabata et al., 1992) embryonic stage 6 -- 17   organism | segment polarity expression pattern   (Tabata et al., 1992) embryonic stage 7 -- 9   organism | segment polarity expression pattern   (Florence et al., 1997) embryonic stage 13 -- 14   large intestine primordium | restricted   (Iwaki et al., 2001) Comment:along outer curve of large intestine third instar larval stage   wing disc | restricted   (Masucci et al., 1990) northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage -- pupal stage     (Drees et al., 1987) embryonic stage 7 -- 15     (Drees et al., 1987) Comment:reference states 3-12 hr AEL
Additional Descriptive Data
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
No Assay Recorded Stage Tissue/Position (including subcellular localization) Reference embryonic stage   ectoderm   (Peifer et al., 1991) embryonic stage 8 -- 11   neuroblast NB7-4   (Doe, 1992) embryonic stage 8 -- 17   intercalary segment   (Schmidt-Ott and Technau, 1992) head   (Schmidt-Ott and Technau, 1992) embryonic stage 9 -- 11   neuroblast NB6-2   (Doe, 1992) embryonic stage 10 -- 11   neuroblast NB6-1   (Doe, 1992) immunolocalization Stage Tissue/Position (including subcellular localization) Reference embryonic stage   organism | segmentally repeated   (DiNardo et al., 1985)   (Manoukian and Krause, 1992) embryonic stage 6   embryonic maxillary segment | posterior   (DiNardo et al., 1985) embryonic stage 7   embryonic maxillary segment | posterior & embryonic labial segment | posterior & embryonic thoracic segment | posteriorembryonic abdominal segment 1 -- embryonic abdominal segment 7   (DiNardo et al., 1985) parasegment 2 -- parasegment 15   (Mlodzik et al., 1990) embryonic stage 7 -- 11   parasegment | restricted & segmentally repeated   (Gonzalez et al., 1991) embryonic stage 8   embryonic gnathal segment | posterior   (DiNardo et al., 1985) embryonic thoracic segment | posterior   (DiNardo et al., 1985) embryonic abdominal segment 1 -- embryonic abdominal segment 9   (DiNardo et al., 1985) embryonic stage 8 -- 11   neuroblast NB7-1   (Doe, 1992) embryonic stage 8 -- 13   organism | segmentally repeated   (Chanas et al., 2004) embryonic stage 8 -- 17   antennal segment   (Schmidt-Ott and Technau, 1992) labral segment   (Schmidt-Ott and Technau, 1992) embryonic stage 9 -- 11   neuroblast NB1-1   (Doe, 1992) neuroblast NB7-2   (Doe, 1992) embryonic stage 10   endoderm | segment polarity expression pattern   (Belenkaya et al., 2002) embryonic stage 10 -- 11   ventral neurogenic region   (Serrano et al., 1995) ventral epidermis primordium   (Serrano et al., 1995) embryonic stage 11   organism | posterior   (Patel et al., 1989) ventral nervous system | posterior & segmentally repeated   (DiNardo et al., 1985) organism | pair rule expression pattern   (Alonso et al., 2001) organism | segment polarity expression pattern   (Hamada and Bienz, 2002, Alonso et al., 2001) embryonic segment | posterior compartment   (Kaminker et al., 2001) organism | segmentally repeated   (Muller and Bienz, 1992) embryonic stage 11 -- 14   organism | segmentally repeated   • nucleus (Larsen et al., 2003) organism | anterior to & segment boundary   • nucleus (Larsen et al., 2003) embryonic stage 11 -- 16   dorsal multidendritic neuron | subset   (Brewster et al., 2001) Comment:expressed in one dorsal multidendritic neuron per segment embryonic stage 12 -- 15   epidermis | segmentally repeated   (Namba et al., 1997) central nervous system | segmentally repeated   (Namba et al., 1997) embryonic stage 13   neuroblast | lateral   (Chang et al., 2000) embryonic stage 13 -- 14   neuromere | segment boundary   (Hirth et al., 2003) leading edge mixer cell | subset   (Gettings et al., 2010) embryonic stage 16   abdominal dorsal multidendritic neuron ddaE | subset   • nucleus (Brewster et al., 2001) Comment:reference states dda1 embryonic stage 17   EW1 neuron   (Lundell and Hirsh, 1998) EW2 neuron   (Lundell and Hirsh, 1998) EW3 neuron   (Lundell and Hirsh, 1998) ventro-lateral serotonergic neuron   (Lundell et al., 1996) third instar larval stage   female genital disc | restricted   (Freeland and Kuhn, 1996) wing disc | posterior compartment   (Chanas et al., 2004) male genital disc | restricted   (Freeland and Kuhn, 1996) in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage 11   organism | segment polarity expression pattern   (Bahri et al., 2001)
Additional Descriptive Data
	The en protein is localized to the nucleus, but becomes diffuse during mitosis. (Patel et al., 1989, DiNardo et al., 1985) The en protein is expressed in the posterior half of each segment. (DiNardo et al., 1985) en protein is expressed in a two cell wide row of cells anterior to the segment boundaries of each segment. Expression in the posterior row of cells decreases as a groove forms at the segment boundary. (Larsen et al., 2003) en expression was observed sequentially in 5 "centers" anterior to the mandibular segment starting at embryonic stage 8. These are the "en antennal stripe", the "en head spot", the "en intercalary spot", the "en expression in the anterior dorsal hemispheres" and the "en expression in the clypeolabrum". Later two of the spots split, generating the "en antennal spot" and the "en secondary head spot". The migration of these en-expressing cells was followed throughout embryonic development. (Schmidt-Ott and Technau, 1992) en protein is expressed in stripes in the posterior region of each segment in the developing embryo. Detection of en expression in the even-numbered stripes slightly preceeds detection in odd-numbered stripes. (Patel et al., 1989) en protein is expressed in 14 stripes beginning at embryonic stage 8. After stage 11, expression does not overlap with that of ci. en protein expression in the wing disc also does not overlap with that of ci (Chanas et al., 2004) The en protein is expressed in a specified subset of neuroblasts in embryonic stages 8-11. (see also FBrf 42042) (Doe, 1992) en expression is diminished in wg and arm mutants. An allelic series was described for each with with increasingly severe segment polarity phenotypes and earlier and more severe loss of en expression. The segmental stripes decay with a similar asymmetric pattern in wg and arm mutants. (Peifer et al., 1991) In wghs.P embryos, the en protein stripes broaden posteriorly, spanning maximally about one-third of the segment. This is a similar expression pattern to that found in nkd mutant embryos. (Noordermeer et al., 1992) Delayed changes in en expression were observed in response to ectopic eve expression in evehs.PS embryos. Four different expression patterns were observed depending on the timing of ectopic eve protein induction. (Manoukian and Krause, 1992)
Marker for
	segment boundary embryonic large intestine
Subcellular Localization
CV Term	nucleus (DiNardo et al., 1985)
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 FBgn0000577    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0000577 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 moderately high expression to a trough of very low expression. Peak expression observed within 00-12 hour embryonic 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 FBgn0000577 expression level Developmental Stage   Expression Level embryo 00-02hr    71 embryo 02-04hr    892 embryo 04-06hr    2966 embryo 06-08hr    3333 embryo 08-10hr    2373 embryo 10-12hr    1327 embryo 12-14hr    887 embryo 14-16hr    428 embryo 16-18hr    248 embryo 18-20hr    290 embryo 20-22hr    244 embryo 22-24hr    203 larva L1    205 larva L2    96 larva L3 12hr old    72 larva L3 puffstage 1-2    123 larva L3 puffstage 3-6    309 larva L3 puffstage 7-9    402 white prepupae new    465 white prepupae 12hr    413 white prepupae 24hr    645 pupae 2d postWPP    312 pupae 3d postWPP    116 pupae 4d postWPP    92 adult male 01day    118 adult male 05day    202 adult male 30day    201 adult female 01day    30 adult female 05day    14 adult female 30day    12 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    71 embryo 02-04hr  (892) embryo 04-06hr  (2966) embryo 06-08hr  (3333) embryo 08-10hr  (2373) embryo 10-12hr  (1327) embryo 12-14hr  (887) embryo 14-16hr  (428) embryo 16-18hr  (248) embryo 18-20hr  (290) embryo 20-22hr  (244) embryo 22-24hr  (203) larva L1  (205) larva L2    96 larva L3 12hr old    72 larva L3 puffstage 1-2  (123) larva L3 puffstage 3-6  (309) larva L3 puffstage 7-9  (402) white prepupae new  (465) white prepupae 12hr  (413) white prepupae 24hr  (645) pupae 2d postWPP  (312) pupae 3d postWPP  (116) pupae 4d postWPP    92 adult male 01day  (118) adult male 05day  (202) adult male 30day  (201) adult female 01day    30 adult female 05day    14 adult female 30day    12 Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    71 embryo 02-04hr    892 embryo 04-06hr  (2966) embryo 06-08hr  (3333) embryo 08-10hr  (2373) embryo 10-12hr    1327 embryo 12-14hr    887 embryo 14-16hr    428 embryo 16-18hr    248 embryo 18-20hr    290 embryo 20-22hr    244 embryo 22-24hr    203 larva L1    205 larva L2    96 larva L3 12hr old    72 larva L3 puffstage 1-2    123 larva L3 puffstage 3-6    309 larva L3 puffstage 7-9    402 white prepupae new    465 white prepupae 12hr    413 white prepupae 24hr    645 pupae 2d postWPP    312 pupae 3d postWPP    116 pupae 4d postWPP    92 adult male 01day    118 adult male 05day    202 adult male 30day    201 adult female 01day    30 adult female 05day    14 adult female 30day    12 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    71 embryo 02-04hr    892 embryo 04-06hr    2966 embryo 06-08hr    3333 embryo 08-10hr    2373 embryo 10-12hr    1327 embryo 12-14hr    887 embryo 14-16hr    428 embryo 16-18hr    248 embryo 18-20hr    290 embryo 20-22hr    244 embryo 22-24hr    203 larva L1    205 larva L2    96 larva L3 12hr old    72 larva L3 puffstage 1-2    123 larva L3 puffstage 3-6    309 larva L3 puffstage 7-9    402 white prepupae new    465 white prepupae 12hr    413 white prepupae 24hr    645 pupae 2d postWPP    312 pupae 3d postWPP    116 pupae 4d postWPP    92 adult male 01day    118 adult male 05day    202 adult male 30day    201 adult female 01day    30 adult female 05day    14 adult female 30day    12 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    71 embryo 02-04hr    892 embryo 04-06hr    2966 embryo 06-08hr    3333 embryo 08-10hr    2373 embryo 10-12hr    1327 embryo 12-14hr    887 embryo 14-16hr    428 embryo 16-18hr    248 embryo 18-20hr    290 embryo 20-22hr    244 embryo 22-24hr    203 larva L1    205 larva L2    96 larva L3 12hr old    72 larva L3 puffstage 1-2    123 larva L3 puffstage 3-6    309 larva L3 puffstage 7-9    402 white prepupae new    465 white prepupae 12hr    413 white prepupae 24hr    645 pupae 2d postWPP    312 pupae 3d postWPP    116 pupae 4d postWPP    92 adult male 01day    118 adult male 05day    202 adult male 30day    201 adult female 01day    30 adult female 05day    14 adult female 30day    12 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0000577 expression level Developmental Stage   Expression Level embryo 00-02hr    71 embryo 02-04hr    892 embryo 04-06hr    2966 embryo 06-08hr    3333 embryo 08-10hr    2373 embryo 10-12hr    1327 embryo 12-14hr    887 embryo 14-16hr    428 embryo 16-18hr    248 embryo 18-20hr    290 embryo 20-22hr    244 embryo 22-24hr    203 larva L1    205 larva L2    96 larva L3 12hr old    72 larva L3 puffstage 1-2    123 larva L3 puffstage 3-6    309 larva L3 puffstage 7-9    402 white prepupae new    465 white prepupae 12hr    413 white prepupae 24hr    645 pupae 2d postWPP    312 pupae 3d postWPP    116 pupae 4d postWPP    92 adult male 01day    118 adult male 05day    202 adult male 30day    201 adult female 01day    30 adult female 05day    14 adult female 30day    12 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    71 embryo 02-04hr  (892) embryo 04-06hr  (2966) embryo 06-08hr  (3333) embryo 08-10hr  (2373) embryo 10-12hr  (1327) embryo 12-14hr  (887) embryo 14-16hr  (428) embryo 16-18hr  (248) embryo 18-20hr  (290) embryo 20-22hr  (244) embryo 22-24hr  (203) larva L1  (205) larva L2    96 larva L3 12hr old    72 larva L3 puffstage 1-2  123 larva L3 puffstage 3-6  (309) larva L3 puffstage 7-9  (402) white prepupae new  (465) white prepupae 12hr  (413) white prepupae 24hr  (645) pupae 2d postWPP  (312) pupae 3d postWPP  116 pupae 4d postWPP    92 adult male 01day  118 adult male 05day  (202) adult male 30day  (201) adult female 01day    30 adult female 05day    14 adult female 30day    12 Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    71 embryo 02-04hr    892 embryo 04-06hr  (2966) embryo 06-08hr  (3333) embryo 08-10hr  2373 embryo 10-12hr    1327 embryo 12-14hr    887 embryo 14-16hr    428 embryo 16-18hr    248 embryo 18-20hr    290 embryo 20-22hr    244 embryo 22-24hr    203 larva L1    205 larva L2    96 larva L3 12hr old    72 larva L3 puffstage 1-2    123 larva L3 puffstage 3-6    309 larva L3 puffstage 7-9    402 white prepupae new    465 white prepupae 12hr    413 white prepupae 24hr    645 pupae 2d postWPP    312 pupae 3d postWPP    116 pupae 4d postWPP    92 adult male 01day    118 adult male 05day    202 adult male 30day    201 adult female 01day    30 adult female 05day    14 adult female 30day    12 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    71 embryo 02-04hr    892 embryo 04-06hr    2966 embryo 06-08hr    3333 embryo 08-10hr    2373 embryo 10-12hr    1327 embryo 12-14hr    887 embryo 14-16hr    428 embryo 16-18hr    248 embryo 18-20hr    290 embryo 20-22hr    244 embryo 22-24hr    203 larva L1    205 larva L2    96 larva L3 12hr old    72 larva L3 puffstage 1-2    123 larva L3 puffstage 3-6    309 larva L3 puffstage 7-9    402 white prepupae new    465 white prepupae 12hr    413 white prepupae 24hr    645 pupae 2d postWPP    312 pupae 3d postWPP    116 pupae 4d postWPP    92 adult male 01day    118 adult male 05day    202 adult male 30day    201 adult female 01day    30 adult female 05day    14 adult female 30day    12 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    71 embryo 02-04hr    892 embryo 04-06hr    2966 embryo 06-08hr    3333 embryo 08-10hr    2373 embryo 10-12hr    1327 embryo 12-14hr    887 embryo 14-16hr    428 embryo 16-18hr    248 embryo 18-20hr    290 embryo 20-22hr    244 embryo 22-24hr    203 larva L1    205 larva L2    96 larva L3 12hr old    72 larva L3 puffstage 1-2    123 larva L3 puffstage 3-6    309 larva L3 puffstage 7-9    402 white prepupae new    465 white prepupae 12hr    413 white prepupae 24hr    645 pupae 2d postWPP    312 pupae 3d postWPP    116 pupae 4d postWPP    92 adult male 01day    118 adult male 05day    202 adult male 30day    201 adult female 01day    30 adult female 05day    14 adult female 30day    12 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 FBgn0000577    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0000577 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: larval hindgut. [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) Linear, scaled to maximum FBgn0000577 expression level Tissue   Expression Level Larval Central Nervous System    46.475 Larval Midgut    3.5 Larval Hindgut    234.3 Larval Malpighian Tubules    0.9 Larval Fat Body    9.9 Larval Salivary Gland    3 Larval Trachea    16.175 Larval Carcass    36.25 Adult Head    21.9 Adult Eye    39.45 Adult Brain    14.5 Adult Thoracic-Abdominal Ganglion    43.8 Adult Crop    7.2 Adult Midgut    0.9 Adult Hindgut    4 Adult Malpighian Tubules    6.1 Adult Fat Body    13.9 Adult Salivary Gland no informative data Adult Heart    2.35 Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data Adult Ovary    2.5 Adult Testis    3.6 Adult Male Accessory Gland    7.3 Adult Carcass    6.7 Expression Level Scale  None   Low   Moderate  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    46.475 Larval Midgut    3.5 Larval Hindgut    234.3 Larval Malpighian Tubules    0.9 Larval Fat Body    9.9 Larval Salivary Gland    3 Larval Trachea    16.175 Larval Carcass    36.25 Adult Head    21.9 Adult Eye    39.45 Adult Brain    14.5 Adult Thoracic-Abdominal Ganglion    43.8 Adult Crop    7.2 Adult Midgut    0.9 Adult Hindgut    4 Adult Malpighian Tubules    6.1 Adult Fat Body    13.9 Adult Salivary Gland no informative data Adult Heart    2.35 Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data Adult Ovary    2.5 Adult Testis    3.6 Adult Male Accessory Gland    7.3 Adult Carcass    6.7 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    46.475 Larval Midgut    3.5 Larval Hindgut    234.3 Larval Malpighian Tubules    0.9 Larval Fat Body    9.9 Larval Salivary Gland    3 Larval Trachea    16.175 Larval Carcass    36.25 Adult Head    21.9 Adult Eye    39.45 Adult Brain    14.5 Adult Thoracic-Abdominal Ganglion    43.8 Adult Crop    7.2 Adult Midgut    0.9 Adult Hindgut    4 Adult Malpighian Tubules    6.1 Adult Fat Body    13.9 Adult Salivary Gland no informative data Adult Heart    2.35 Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data Adult Ovary    2.5 Adult Testis    3.6 Adult Male Accessory Gland    7.3 Adult Carcass    6.7 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    46.475 Larval Midgut    3.5 Larval Hindgut    234.3 Larval Malpighian Tubules    0.9 Larval Fat Body    9.9 Larval Salivary Gland    3 Larval Trachea    16.175 Larval Carcass    36.25 Adult Head    21.9 Adult Eye    39.45 Adult Brain    14.5 Adult Thoracic-Abdominal Ganglion    43.8 Adult Crop    7.2 Adult Midgut    0.9 Adult Hindgut    4 Adult Malpighian Tubules    6.1 Adult Fat Body    13.9 Adult Salivary Gland no informative data Adult Heart    2.35 Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data Adult Ovary    2.5 Adult Testis    3.6 Adult Male Accessory Gland    7.3 Adult Carcass    6.7 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to maximum FBgn0000577 expression level Tissue   Expression Level Larval Central Nervous System    46.475 Larval Midgut    3.5 Larval Hindgut    234.3 Larval Malpighian Tubules    0.9 Larval Fat Body    9.9 Larval Salivary Gland    3 Larval Trachea    16.175 Larval Carcass    36.25 Adult Head    21.9 Adult Eye    39.45 Adult Brain    14.5 Adult Thoracic-Abdominal Ganglion    43.8 Adult Crop    7.2 Adult Midgut    0.9 Adult Hindgut    4 Adult Malpighian Tubules    6.1 Adult Fat Body    13.9 Adult Salivary Gland no informative data Adult Heart    2.35 Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data Adult Ovary    2.5 Adult Testis    3.6 Adult Male Accessory Gland    7.3 Adult Carcass    6.7 Expression Level Scale  None   Low   Moderate  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    46.475 Larval Midgut    3.5 Larval Hindgut    234.3 Larval Malpighian Tubules    0.9 Larval Fat Body    9.9 Larval Salivary Gland    3 Larval Trachea    16.175 Larval Carcass    36.25 Adult Head    21.9 Adult Eye    39.45 Adult Brain    14.5 Adult Thoracic-Abdominal Ganglion    43.8 Adult Crop    7.2 Adult Midgut    0.9 Adult Hindgut    4 Adult Malpighian Tubules    6.1 Adult Fat Body    13.9 Adult Salivary Gland no informative data Adult Heart    2.35 Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data Adult Ovary    2.5 Adult Testis    3.6 Adult Male Accessory Gland    7.3 Adult Carcass    6.7 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    46.475 Larval Midgut    3.5 Larval Hindgut    234.3 Larval Malpighian Tubules    0.9 Larval Fat Body    9.9 Larval Salivary Gland    3 Larval Trachea    16.175 Larval Carcass    36.25 Adult Head    21.9 Adult Eye    39.45 Adult Brain    14.5 Adult Thoracic-Abdominal Ganglion    43.8 Adult Crop    7.2 Adult Midgut    0.9 Adult Hindgut    4 Adult Malpighian Tubules    6.1 Adult Fat Body    13.9 Adult Salivary Gland no informative data Adult Heart    2.35 Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data Adult Ovary    2.5 Adult Testis    3.6 Adult Male Accessory Gland    7.3 Adult Carcass    6.7 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    46.475 Larval Midgut    3.5 Larval Hindgut    234.3 Larval Malpighian Tubules    0.9 Larval Fat Body    9.9 Larval Salivary Gland    3 Larval Trachea    16.175 Larval Carcass    36.25 Adult Head    21.9 Adult Eye    39.45 Adult Brain    14.5 Adult Thoracic-Abdominal Ganglion    43.8 Adult Crop    7.2 Adult Midgut    0.9 Adult Hindgut    4 Adult Malpighian Tubules    6.1 Adult Fat Body    13.9 Adult Salivary Gland no informative data Adult Heart    2.35 Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data Adult Ovary    2.5 Adult Testis    3.6 Adult Male Accessory Gland    7.3 Adult Carcass    6.7 Expression Level Scale  None   Low   Moderate   High   Very high  Heatmap Tissue   Expression Level Larval Central Nervous System     Larval Midgut     Larval Hindgut     Larval Malpighian Tubules     Larval Fat Body     Larval Salivary Gland     Larval Trachea     Larval Carcass     Adult Head     Adult Eye     Adult Brain     Adult Thoracic-Abdominal Ganglion     Adult Crop     Adult Midgut     Adult Hindgut     Adult Malpighian Tubules     Adult Fat Body     Adult Salivary Gland no informative data Adult Heart     Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data Adult Ovary     Adult Testis     Adult Male Accessory Gland     Adult Carcass     FlyAtlas Organ/Tissue Expression, larval vs. adult Larval Expression Level Tissue Adult Expression Level   NA  Head    21.9   NA  Eye    39.45   NA  Brain    14.5   46.475  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    43.8   NA  Crop    7.2   3.5  Midgut    0.9   234.3  Hindgut    4   0.9  Malpighian Tubules    6.1   9.9  Fat Body    13.9   3  Salivary Gland    no informative data   NA  Heart    2.35   16.175  Trachea    NA   NA  VirginFemale Spermatheca    no informative data   NA  InseminatedFemale Spermatheca    no informative data   NA  Ovary    2.5   NA  Testis    3.6   NA  Male Accessory Gland    7.3   36.25  Carcass    6.7 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_11
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0000577 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG9015-RB FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for en Stages(s) 4-6 Stages(s) 7-8 Stages(s) 9-10 Stages(s) 11-12 Stages(s) 13-16
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal en1 en2 en3 en4 en8 en30 en31 en32 en34 en37 en38 en43 en47 en48 en49 en50 en51 en52 en53 en57 en58 en59 enSF59 lethal (with Df(2R)enE) enAdv-1 lethal (with Df(2R)enF) enAdv-1 lethal (with Df(2R)en-A) enAdv-1 lethal (with Df(2R)en-B) enAdv-1 lethal (with Df(2R)en-SFX31) enAdv-1 lethal (with en54) enAdv-1 lethal (with en57) enAdv-1 lethal (with en59) enAdv-1 lethal (with enAdv-1) en54 en57 en59 enXho25 lethal (with enXho25) enAdv-1 lethal | embryonic stage, with Scer\GAL4arm.PS enScer\UAS.cGa lethal | embryonic stage | pair rule expression pattern | recessive enunspecified lethal | embryonic stage | recessive en2 en4 en6 en7 en10 en34 en47 en48 en49 en50 en54 en57 en58 en63 enCX1 enE ennst-101 ennst-120 lethal | embryonic stage | segment polarity expression pattern, with Scer\GAL4nos.PG enScer\UAS.cYa lethal | heat sensitive enhs.3 lethal | recessive en2 en3 en4 en5 en6 en7 en8 en9 en10 en12 en13 en14 en15 en16 en17 en18 en19 en20 en21 en22 en23 en24 en25 en26 en27 en28 en29 en30 en31 en32 en33 en34 en35 en36 en37 en38 en39 en40 en41 en42 en43 en44 en45 en46 en49 en59 enAdv-1 enC enEnci enLI034 enSFX22 enWR1 enWR3 enXho25 semi-lethal | pupal stage, with Scer\GAL4pnr-MD237 enGD13214 viable en1 en10B-5 enF4 viable (with Df(2R)en30) enAdv-1 viable (with Df(2R)enC) enAdv-1 viable (with en1) enAdv-1 viable (with enAdv-1) en1 Sterility Allele fertile | female | reduced enspt sterile en10B-5 Other Phenotypes Allele neuroanatomy defective en54 endsRNA.cIa neuroanatomy defective, with Scer\GAL4sim.P3.7 enScer\UAS.cGa neuroanatomy defective, with Scer\GAL4sim.PS enScer\UAS.cYa neuroanatomy defective | embryonic stage enC planar polarity defective, with Scer\GAL469B enScer\UAS.cGa planar polarity defective, with Scer\GAL4Ubx.PdC enScer\UAS.cGa visible en1 en59 enF4 enF enαTub84B.PZ visible, with Scer\GAL469B enScer\UAS.cGa visible, with Scer\GAL4ey.PH enScer\UAS.cYa visible, with Scer\GAL4pnr-MD237 enGD13214 visible, with Scer\GAL4T155 enScer\UAS.cGa visible, with Scer\GAL4Ubx.PdC enScer\UAS.cGa visible (with Df(2R)en30) enAdv-1 visible (with Df(2R)enC) enAdv-1 visible (with en1) enAdv-1 visible (with enAdv-1) en1 visible | dominant enApa enEs visible | heat sensitive enspt visible | recessive en1 en54 enWR1 enWR3 Phenotype manifest in Allele abdominal 1 ventral denticle belt en4 en57 en58 en6 abdominal 2 ventral denticle belt en4 en57 en58 en6 abdominal 3 ventral denticle belt en4 en57 en58 en6 abdominal 4 ventral denticle belt en4 en57 en58 en6 abdominal 5 ventral denticle belt en4 en57 en58 en6 abdominal 6 ventral denticle belt en4 en57 en58 en6 abdominal 7 ventral denticle belt en4 en57 en58 en6 abdominal 8 ventral denticle belt en57 en58 abdominal segment 6 | pair rule expression pattern enunspecified abdominal segment 8 | pair rule expression pattern enunspecified abdominal tergite & macrochaeta enApa enEs abdominal tergite & trichome enApa enEs abdominal tergite | anterior & trichome enαTub84B.PZ adult abdomen enEs adult head, with Scer\GAL4ey.PH enScer\UAS.cYa alar lobe en1 anal plate | somatic clone en57 en59 antenna en1 antennal sense organ enCX1 anterior commissure en54 anterior commissure, with Scer\GAL4elav-C155 enScer\UAS.cYa anterior commissure, with Scer\GAL4sim.PS enScer\UAS.cYa anterior crossvein en1 apodeme | embryonic stage en54 basal apodeme of penis enαTub84B.PZ central nervous system | somatic clone en57 en59 cephalopharyngeal skeleton | heat sensitive enhs.3 clasper en1 en2 en3 en4 en6 en7 clasper | somatic clone en57 en59 commissure | embryonic stage enC enE commissure | embryonic stage, with Scer\GAL4elav-C155 enScer\UAS.cYa connective, with Scer\GAL4elav-C155 enScer\UAS.cYa costal cell, with Scer\GAL430A enScer\UAS.cGa cuticle en4 en54 denticle belt en2 en4 en54 en59 en7 enC enCX1 endsRNA.cMa enhs.PGa enhs.PH denticle belt | heat sensitive enhs.3 denticle belt | somatic clone en4 dorsal triple row, with Scer\GAL4112-97 enScer\UAS.cTa dorsal triple row, with Scer\GAL4en-e16E enScer\UAS.cTa EL neuron enhs.3 embryo, with Scer\GAL4arm.PS enScer\UAS.cGa embryo | posterior compartment en59 embryo | segment polarity expression pattern, with Scer\GAL4nos.PG enScer\UAS.cYa embryonic/first instar larval cuticle en54 en58 enC enhs.PH embryonic/first instar larval cuticle, with Scer\GAL4arm.PS enScer\UAS.cGa embryonic/first instar larval cuticle, with Scer\GAL4prd.RG1 enScer\UAS.cTa embryonic/larval central nervous system enC embryonic/larval cuticle en59 embryonic/larval fat body en4 embryonic/larval heart, with Scer\GAL4twi.PGa enScer\UAS.cGa embryonic/larval pericardial cell & embryonic myoblast, with Scer\GAL4how-24B, Scer\GAL4twi.PG enScer\UAS.cGa embryonic/larval peripheral nervous system enC embryonic/larval somatic muscle en54 embryonic/larval somatic muscle, with Scer\GAL4how-24B, Scer\GAL4twi.PG enScer\UAS.cGa embryonic/larval somatic muscle, with Scer\GAL4twi.PGa enScer\UAS.cGa embryonic abdomen en54 en58 embryonic abdominal segment 2 | pair rule expression pattern enunspecified embryonic abdominal segment 4 | pair rule expression pattern enunspecified embryonic epidermis en9.6 embryonic epidermis | heat sensitive enhs.3 embryonic gnathal segment en4 embryonic head en54 en58 embryonic head | heat sensitive enhs.3 embryonic hindgut, with Scer\GAL4byn-Gal4 enScer\UAS.cTa enScer\UAS.cYa embryonic leading edge cell, with Scer\GAL4ptc-559.1 enGD13214 embryonic metathoracic segment | pair rule expression pattern enunspecified embryonic myoblast, with Scer\GAL4how-24B, Scer\GAL4twi.PG enScer\UAS.cGa embryonic prothoracic segment | pair rule expression pattern enunspecified embryonic segment enLI034 epidermis | posterior | somatic clone en57 en59 eye, with Scer\GAL4ey.PH enScer\UAS.cYa fascicle, with Scer\GAL4elav-C155 enScer\UAS.cYa fat body/gonad primordium & parasegment 4, with Scer\GAL4twi.PGa enScer\UAS.cGa fat body/gonad primordium & parasegment 5, with Scer\GAL4twi.PGa enScer\UAS.cGa fat body/gonad primordium & parasegment 6, with Scer\GAL4twi.PGa enScer\UAS.cGa fat body/gonad primordium & parasegment 6, with Scer\GAL4zen.Kr.PF enScer\UAS.cGa fat body/gonad primordium & parasegment 7, with Scer\GAL4twi.PGa enScer\UAS.cGa fat body/gonad primordium & parasegment 7, with Scer\GAL4zen.Kr.PF enScer\UAS.cGa fat body/gonad primordium & parasegment 8, with Scer\GAL4twi.PGa enScer\UAS.cGa fat body/gonad primordium & parasegment 8, with Scer\GAL4zen.Kr.PF enScer\UAS.cGa fat body/gonad primordium & parasegment 9, with Scer\GAL4twi.PGa enScer\UAS.cGa female gonopod en1 en2 genital arch en1 en2 en3 en4 en6 en7 genital disc primordium enE gonad en4 en56 gonad | embryonic stage (with en22.48) en60.04 gonad | embryonic stage (with en60.04) en22.48 gonadal sheath proper primordium | embryonic stage (with en22.48) en60.04 gonadal sheath proper primordium | embryonic stage (with en60.04) en22.48 gonopod long bristle enαTub84B.PZ gonopod thorn bristle enαTub84B.PZ haltere, with Scer\GAL4112-97 enScer\UAS.cTa haltere | anterior compartment enαTub84B.PZ hypandrial bristle enαTub84B.PZ hypandrial process enαTub84B.PZ hypandrium en1 en2 en3 en4 en6 en7 hypandrium | somatic clone en57 en59 imaginal disc | posterior compartment en59 intersegmental nerve, with Scer\GAL4elav-C155 enScer\UAS.cYa Keilin's organ en10 en57 en59 en7 enX49 lateral plate | somatic clone en57 en59 lateral tract en54 leg | posterior | somatic clone en57 en59 longitudinal connective enC enE longitudinal connective, with Scer\GAL4elav-C155 enScer\UAS.cYa male genitalia enαTub84B.PZ male genitalia | cell non-autonomous | somatic clone en59 male genitalia | somatic clone en57 male terminalia sensillum enαTub84B.PZ medial triple row, with Scer\GAL4112-97 enScer\UAS.cTa medial triple row, with Scer\GAL4en-e16E enScer\UAS.cTa mesothoracic tarsal bristle longitudinal row 2 en1 mesothoracic tarsal bristle longitudinal row 2.5 en1 mesothoracic tergum, with Scer\GAL4pnr-MD237 enGD13214 mesothoracic tibial transverse bristle row en1 metathoracic tibial transverse bristle row en1 metathoracic ventral denticle belt en4 en6 midline glial cell, with Scer\GAL4sim.P3.7 enScer\UAS.cGa midline glial cell, with Scer\GAL4sim.PS enScer\UAS.cYa midline neuronal primordium, with Scer\GAL4sim.PS enScer\UAS.cYa motor neuron & growth cone & embryo, with Scer\GAL4elav-C155 enScer\UAS.cYa MP1 neuron, with Scer\GAL4elav-C155 enScer\UAS.cYa MP1 neuron, with Scer\GAL4sim.PS enScer\UAS.cYa neuroblast NB7-3 | ectopic enhs.3 neuromuscular junction | embryonic stage, with Scer\GAL4elav-C155 enScer\UAS.cYa penis enαTub84B.PZ en1 en2 en3 en4 en6 en7 posterior commissure en54 posterior commissure, with Scer\GAL4elav-C155 enScer\UAS.cYa posterior commissure, with Scer\GAL4sim.PS enScer\UAS.cYa presumptive embryonic/larval central nervous system enhs.3 presumptive embryonic/larval central nervous system, with Scer\GAL4elav-C155 enScer\UAS.cYa presumptive embryonic/larval muscle system enC enE presumptive embryonic/larval peripheral nervous system, with Scer\GAL4elav-C155 enScer\UAS.cYa proboscis | posterior | somatic clone en57 en59 prothoracic leg en1 prothoracic tarsal segment | posterior compartment en1 RP neuron, with Scer\GAL4elav-C155 enScer\UAS.cYa scutellum en1 scutellum (with en1) enAdv-1 scutellum (with enAdv-1) en1 second segment of antenna | posterior compartment (with en1) en2 second segment of antenna | posterior compartment (with en2) en1 segment | posterior compartment en54 segmental nerve, with Scer\GAL4elav-C155 enScer\UAS.cYa sensillum campaniformium en1 sensillum campaniformium of anterior crossvein en1 serotonergic neuron | embryonic stage enC sex comb en1 enEnci sex comb | ectopic en1 spermathecal duct enspt spermathecum enspt spermathecum | heat sensitive enspt stigmatophore enE tarsal segment enCX1 tergite enαTub84B.PZ enApa enEs tergite, with Scer\GAL4T155 enScer\UAS.cGa tergite | anterior compartment enApa tergite | anterior compartment, with Scer\GAL469B enScer\UAS.cGa tergite | anterior compartment, with Scer\GAL4T155 enScer\UAS.cGa tergite | anterior compartment, with Scer\GAL4Ubx.PdC enScer\UAS.cGa tergite | ectopic | posterior compartment enApa tergite | ectopic | posterior compartment, with Scer\GAL4T155 enScer\UAS.cGa tergite | posterior compartment, with Scer\GAL4Ubx.PdC enScer\UAS.cGa terminal filament | somatic clone enE third segment of antenna en1 trichome of the posterior wing margin en1 enSF64 enSF65 enSF66 UMI interneuron & axon, with Scer\GAL4sim.PS enScer\UAS.cYa ventral nerve cord en54 ventral triple row, with Scer\GAL4112-97 enScer\UAS.cTa ventral triple row, with Scer\GAL4en-e16E enScer\UAS.cTa visceral mesoderm | precursor, with Scer\GAL4twi.PGa enScer\UAS.cGa VUM neuron & axon, with Scer\GAL4sim.PS enScer\UAS.cYa wing en1 en4 en7 en9.6 enC enCX1 enE enF4 wing, with Scer\GAL4112-97 enScer\UAS.cTa wing (with Df(2R)en30) enAdv-1 wing (with Df(2R)enC) enAdv-1 wing (with en1) en28 enAdv-1 wing (with en28) en1 wing (with en54) enAdv-1 wing (with enAdv-1) en1 en54 wing & macrochaeta en1 enEnci wing | anterior/posterior compartment boundary enE wing | anterior/posterior compartment boundary | somatic clone en57 en59 wing | anterior | proximal, with Scer\GAL430A enScer\UAS.cSa wing | anterior | proximal, with Scer\GAL4vg.PM enScer\UAS.cSa wing | anterior compartment, with Scer\GAL4vg.PM enScer\UAS.cSa wing | anterior compartment | dorsal, with Scer\GAL4Bx-MS1096 enScer\UAS.cGa wing | anterior compartment | somatic clone enαTub84B.PZ wing | dorsal | posterior compartment, with Scer\GAL4Bx-MS1096 enScer\UAS.cGa wing | posterior en1 wing | posterior | somatic clone en57 en59 wing | posterior compartment en1 en8 enEnci enEs wing | posterior compartment, with Scer\GAL4en-e16E enScer\UAS.cTa wing | posterior compartment, with Scer\GAL4vg.PM enScer\UAS.cSa wing | posterior compartment | somatic clone en1 wing | somatic clone en4 wing disc | anterior/posterior compartment boundary | somatic clone en1 wing disc | anterior | somatic clone enαTub84B.PZ wing disc | posterior compartment | somatic clone en1 wing vein enαTub84B.PZ en1 enE enEnci enSF64 enSF65 enSF66 enSac27 enXho23 wing vein, with Scer\GAL4112-97 enScer\UAS.cTa wing vein, with Scer\GAL4en-e16E enScer\UAS.cTa wing vein | anterior compartment, with Scer\GAL471B enScer\UAS.cGa wing vein | anterior compartment, with Scer\GAL4C-734 enScer\UAS.cGa wing vein L3 en1 wing vein L4 en1 en54 en59 wing vein L5 en1 enF4
Classical Alleles ( 113 )
For All Classical Alleles Show
Allele of en Class Mutagen Stocks Known lesion en1 loss of function allele spontaneous 9 Yes en58 loss of function allele ethyl methanesulfonate 7 Yes en54 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate 2 Yes en4 ethyl methanesulfonate 2 Yes en7 ethyl methanesulfonate 2 Yes enApa gain of function allele 2 Yes enXho25 P-element activity p pi25.7wc 2 Yes en59 ethyl methanesulfonate 1 Yes enAuI Delta2-3 1 -- enE Delta2-3 1 Yes enspt hypomorphic allele - genetic evidence spontaneous 1 -- en8 amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes en9.6 amorphic allele - genetic evidence ethyl methanesulfonate 0 -- en96 amorphic allele - genetic evidence gamma ray 0 Yes enCX1 amorphic allele - genetic evidence, loss of function allele 0 Yes enLI034 amorphic allele - genetic evidence 0 -- en+I.CyO wild-type allele natural population 0 Yes en+I.OR wild-type allele natural population 0 -- en+I.Uk wild-type allele natural population 0 Yes en10 ethyl methanesulfonate 0 -- en10B-5 Delta2-3 0 Yes en12 ethyl methanesulfonate 0 -- en12A-5R loss of function allele Delta2-3 0 Yes en13 ethyl methanesulfonate 0 -- en14 ethyl methanesulfonate 0 -- en15 ethyl methanesulfonate 0 -- en16 ethyl methanesulfonate 0 -- en17 gamma ray 0 -- en17en52 P-element activity 0 Yes en18 formaldehyde 0 -- en19 formaldehyde 0 -- en20 formaldehyde 0 -- en21 formaldehyde 0 -- en22.48 ethyl methanesulfonate 0 -- en22 formaldehyde 0 -- en23 formaldehyde 0 -- en24 formaldehyde 0 Yes en25 formaldehyde 0 -- en26 formaldehyde 0 -- en27 formaldehyde 0 -- en28-11-1 0 -- en28 formaldehyde 0 -- en29 formaldehyde 0 -- en2 ethyl methanesulfonate 0 Yes en30 formaldehyde 0 Yes en31 formaldehyde 0 -- en32 X ray 0 -- en33 X ray 0 -- en34 X ray ethyl methanesulfonate 0 Yes en35 X ray 0 -- en36 X ray 0 -- en37 X ray 0 -- en38 X ray 0 Yes en39 X ray 0 -- en3 ethyl methanesulfonate 0 Yes en40 X ray 0 -- en41 X ray 0 -- en42 X ray 0 -- en43 X ray 0 Yes en44 X ray 0 -- en45 X ray 0 -- en46 X ray 0 -- en47 X ray 0 Yes en48 X ray 0 Yes en49 X ray 0 Yes en50 X ray 0 Yes en51 X ray 0 Yes en52 X ray 0 Yes en53 X ray 0 Yes en55 ethyl methanesulfonate 0 -- en56 ethyl methanesulfonate 0 -- en57 ethyl methanesulfonate 0 Yes en5 ethyl methanesulfonate 0 -- en60.04 ethyl methanesulfonate 0 -- en61 0 Yes en63 0 Yes en67 0 Yes en6 ethyl methanesulfonate 0 Yes en9 ethyl methanesulfonate 0 Yes enAdv-1 0 -- enB24 0 -- enC Delta2-3 0 Yes ene15A P-element activity 0 -- ene21A P-element activity 0 -- ene24A P-element activity 0 -- ene33A P-element activity 0 -- ene43A P-element activity 0 -- ene74A P-element activity 0 -- ene76B P-element activity 0 -- enEnci gamma ray 0 -- enEs gain of function allele X ray 0 Yes enF4 P-element activity 0 -- enF hypomorphic allele - genetic evidence Delta2-3 0 Yes enGAL4-33 gene conversion 0 Yes enGAL4-54 gene conversion 0 Yes ennst-101 P-element activity 0 -- ennst-120 P-element activity 0 Yes enSac27 hypomorphic allele - genetic evidence p pi25.7wc 0 Yes enSF59 0 Yes enSF64 X ray 0 Yes enSF65 X ray 0 Yes enSF66 X ray 0 Yes enSF67 0 Yes enSF68 X ray 0 Yes enSF69 X ray 0 Yes enSF7 0 -- enSFX22 X ray 0 -- enT84 0 -- enunspecified 0 -- enWR1 gamma ray 0 Yes enWR3 gamma ray 0 -- enX49 0 -- enXho23 p pi25.7wc 0 Yes
Alleles Carried on Transgenic Constructs ( 71 )
For All Alleles Carried on Transgenic Constructs Show
Allele of en Class Mutagen Stocks Known lesion enGD13214 in vitro construct - RNAi in vitro construct - regulatory fusion 2 Yes enHMS00595 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes enhs.3 in vitro construct - regulatory fusion 1 Yes enJF02316 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes enKK107874 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes enScer\UAS.cYa in vitro construct - regulatory fusion 1 Yes enT:Avic\GFP-EGFP in vitro construct - coding region fusion 1 Yes en31Rg in vitro construct - genomic fragment 0 Yes en406-STOP in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes en81-STOP in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes enABCDe.T:Scer\GAL4 in vitro construct - coding region fusion 0 Yes enABCDEFG in vitro construct - deletion 0 Yes enABCDEFGH in vitro construct 0 Yes enABCDEFH in vitro construct - deletion 0 Yes enABCDEGH in vitro construct - deletion 0 Yes enABCDFGH in vitro construct - deletion 0 Yes enABCDGH in vitro construct - deletion 0 Yes enABCEFGH in vitro construct - deletion 0 Yes enABCGH in vitro construct - deletion 0 Yes enABDEFGH in vitro construct - deletion 0 Yes enABDGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enABGH in vitro construct - deletion 0 Yes enACDEFGH in vitro construct - deletion 0 Yes enACDGH in vitro construct - deletion 0 Yes enACGH in vitro construct - deletion 0 Yes enAct5C.P in vitro construct - regulatory fusion 0 Yes enAct5C.PG in vitro construct - regulatory fusion 0 Yes enAct5C.PH in vitro construct - regulatory fusion 0 Yes enAct5C.PJ in vitro construct - regulatory fusion 0 Yes enADGH in vitro construct - deletion 0 Yes enAGH in vitro construct - deletion 0 Yes enBCDEFGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enBCDEGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enBCDGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enBCGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enBDGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enBGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enCDEFGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enCDEGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enCDGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enCGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enD.T:Scer\GAL4 in vitro construct - coding region fusion 0 Yes enDEFGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enDEGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enDGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes endsRNA.cIa in vitro construct - RNAi 0 Yes endsRNA.cMa in vitro construct - RNAi 0 Yes enEFGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enEGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enF175E in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes enFGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enGH.T:Ivir\HA1 in vitro construct - deletion in vitro construct - coding region fusion 0 Yes enhs.P in vitro construct - regulatory fusion 0 Yes enhs.PGa in vitro construct - regulatory fusion 0 Yes enhs.PGb in vitro construct - regulatory fusion 0 Yes enhs.PH in vitro construct - regulatory fusion 0 Yes enScer\FRT.Rnor\CD2.αTub84B in vitro construct - regulatory fusion 0 Yes enScer\FRT.Scer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes enScer\FRT.αTub84B in vitro construct - regulatory fusion 0 Yes enScer\UAS.cGa in vitro construct - regulatory fusion 0 Yes enScer\UAS.cKa in vitro construct - regulatory fusion 0 Yes enScer\UAS.cSa in vitro construct - regulatory fusion 0 Yes enScer\UAS.cTa in vitro construct - regulatory fusion 0 Yes enScer\UAS.cUa in vitro construct - regulatory fusion 0 Yes enScer\UAS.T:Avic\GFP-m5 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes enScer\UAS.T:Hsim\VP16 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes enScer\UAS.T:Ivir\HA1 FLPase 0 Yes enScer\UAS.Δ282.T:Hsim\VP16 in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - deletion 0 Yes entCH322-92I14 in vitro construct - genomic fragment 0 Yes enαTub84B.PZ FLPase 0 Yes enΔeh1 in vitro construct - deletion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2R)en-11 (van den Heuvel et al., 1993, Sanicola et al., 1995) Df(2R)en28 (Harbecke and Lengyel, 1995, Locke and Hanna, 1996) Df(2R)enC (Saenz-Robles et al., 1995) Df(2R)enD (Saenz-Robles et al., 1995) Df(2R)enE (Lawrence et al., 1999, Sampedro et al., 1993, Cadigan et al., 1994, Hidalgo, 1994, Zecca et al., 1995, Tabata et al., 1995, Gustavson et al., 1996, Martin-Bermudo and Brown, 1996, Matsuzaki and Saigo, 1996, Strutt and Mlodzik, 1996, Azpiazu et al., 1996, Chen and Baker, 1997, Lawrence et al., 1996, Whiteley and Kassis, 1997, Blair and Ralston, 1997, Zhai et al., 2009) Df(2R)enF (Gustavson et al., 1996) Df(2R)en-A (Eberlein and Russell, 1983, Chase and Baker, 1995, Harbecke and Lengyel, 1995, Bhojwani et al., 1995) Df(2R)en-B (Eberlein and Russell, 1983, van den Heuvel et al., 1993, Chase and Baker, 1995, Harbecke and Lengyel, 1995, Poux et al., 1996) Df(2R)en-SFX31 (Karr et al., 1989, Hama et al., 1990, Serrano et al., 1995, Chase and Baker, 1995, Harbecke and Lengyel, 1995, Soto et al., 1995, Bhojwani et al., 1995, Gustavson et al., 1996, Whiteley and Kassis, 1997, Joly et al., 2007) Df(2R)G44 (Muller et al., 1999) Df(2R)I704 (Muller et al., 1999) In(2R)Adv (Srivastava et al., 2002) In(2R)enCX1 (Saenz-Robles et al., 1995) In(2R)enEnci (Locke and Hanna, 1996) In(2R)vgWR1 (Williams et al., 1990) In(2R)vgWR3 (Williams et al., 1990) T(2;3)enEs (Epper and Sanchez, 1983, Kopp et al., 1997) Ts(YLt;2Lt)H144 (Muller et al., 1999)
Duplicated in	Dp(2;3)707   Dp(2;3)en+ (Locke and Hanna, 1996)
Partially disrupted in	Df(2R)en30 (Russell and Eberlein, 1979, Eberlein and Russell, 1983)
Not disrupted in	Df(2R)Np (Bridges et al., 1936) In(2R)vgW (Williams et al., 1990)
Transgenic Constructs & Insertions
Transgenic Constructs
heat-shock construct	Name Expression Data P{enhs.P} NA P{hs-en3} NA P{HS-en.G} NA P{hs-en.H} NA
UAS construct	Name Expression Data P{GD13214} NA P{KK107874} NA P{TRiP.HMS00595} NA P{TRiP.JF02316} NA P{UAS(FRT.y+CD2)flu-en} NA P{UAS(-FRT)flu-en} NA P{UAS-en.G} NA P{UAS-en.S} NA P{UAS-en.T:VP16} NA P{UAS-en.T} NA P{UAS-en.U} NA P{UAS-en.Y} NA P{UAS-en.Δ282::VP16} NA
reporter construct	Name Expression Data P{en1(w)} No P{en1} No P{en/lac} No P{en.96-EZ400} No P{en.96-HZ73(ry)} No P{en.96-HZ73(w)} No P{enB} No P{enC} No P{enD} No P{enE} No P{enF} No P{enG} No P{enH} No P{enH+Hox.A} No P{enH+Hox.B} No P{enI} No P{enJ} No P{en-lacZ.G} No P{en-lacZ.K} No P{en-lacZ(neoSac)} No P{en-lacZ(rySac)} No P{en-lacZ(ryStu)} No P{en-lacZ(ryXba)} No P{en-lacZ(wSac)} No P{en-lacZ(wStu)} No P{en-lacZ(wXba)} No P{en-lacZ(Xho)} No P{en-lacZ} No
characterization construct	Name Expression Data P{2(wen.6)} NA P{3'C-I} NA P{3'wen.6} NA P{3xC} NA P{31Rg-en} NA P{A-I} NA P{A-P} NA P{B-E} NA P{B-F} NA P{B-I.[hd-]} NA P{B-I.[nhr-]} NA P{B-I} NA P{C-I} NA P{D-I} NA P{en2} NA P{en3} NA P{en::Su(H).FRT.T} NA P{en::Su(H).hs.2sev.T} NA P{en.SR} NA P{enDV1} NA P{enDV2} NA P{enF->E} NA P{en-FLP1.L} NA P{enH(w)} NA P{en-w.2.6D} NA P{en-w.2.6DR} NA P{en-w.2.6lacZ} NA P{en-w.2.6up} NA P{en-w.2.6upR} NA P{en-w.13} NA P{en-w.14} NA P{en-w.16-4} NA P{en-w.98bp} NA P{en-w.139bp} NA P{en-w.139GBS} NA P{en-w.181D} NA P{en-w.181lacZ} NA P{en-w.181upR} NA P{en-w.MutGAGA} NA P{en-w.MutNC1} NA P{en-w.MutNC2} NA P{en-w.MutSite1} NA P{en-w.MutSite2} NA P{en-w.PRE} NA P{en-w.Δ1} NA P{en-w.Δ2} NA P{en-w.Δ4} NA P{en-w.Δ5} NA P{enΔeh1} NA P{hb-en::h.J} NA P{hs-EFE} NA P{hs-EFEΔ3} NA P{hs-EFEΔ4} NA P{hs-EFEΔ5} NA P{hs-EFEΔ6'} NA P{hs-EFEΔ6} NA P{hs-EFEΔ23} NA P{hs-EFEΔ34} NA P{hs-EFEΔ46} NA P{hs-EFEΔ234} NA P{hs-EFEΔeh1} NA P{hs-EFEΔF->E} NA P{hs-en::h.J} NA P{hs-Meh1} NA P{mos-en} NA P{N+-G.4,enRep} NA P{ovoD1.EN} NA P{ovoEN} NA P{UAS.ey1-545.enRD} NA P{UAS-ci.ΔEn} NA P{UAS-ciZn/EnRD} NA P{UAS-en::onecut.N} NA P{UAS-en::ovo.B2} NA P{UAS-en::ovo.D1} NA P{UAS-en::sr.rep} NA P{UAS-en-hth.1-430} NA P{UAS-en-Meis3.1-133} NA P{UAS-osaRD} NA P{wen.1} NA P{wen.2} NA P{wen.3} NA P{wen.4} NA P{wen.5} NA P{wen.6} NA P{wen.7} NA P{wen.8.TACG} NA P{wen.8.Δ17} NA P{wen.8} NA P{wen.10} NA P{wen.12} NA P{wen.V8} NA P{αTub84B(FRT.f+)en.Z1} NA P{αTub84B(FRT.y+CD2)en.Z2} NA P{αTub84B(-FRT)en.Z1} NA P{αTub84B(-FRT)en.Z2} NA PBac{CH322-92I14} NA PBac{en-EGFP.S} NA
Insertions
	Type of insertions Name Expression data insertion of enhancer trap P{en1}48A No P{en1}en17en52 No P{lacW}ennst-101 No P{lwB}enAuI No miscellaneous insertions ?{}en1 NA P{enF}enF4 NA P{en-lacZ(neoSac)}enSac27 NA P{en-lacZ(Xho)}enXho23 NA P{en-lacZ(Xho)}enXho25 NA P{mos-en}en10B-5 NA P{mos-en}en12A-5R NA insertion of enhancer trap binary system P{en2.4-GAL4}ene15A No P{en2.4-GAL4}ene21A No P{en2.4-GAL4}ene24A No P{en2.4-GAL4}ene33A No P{en2.4-GAL4}ene43A No P{en2.4-GAL4}ene74A No P{en2.4-GAL4}ene76B No P{GAL4}enGAL4-33 No P{GAL4}enGAL4-54 No
Gene Ontology: Function, Process & Cellular Component ( 29 unique terms )
Terms Based on Experimental Evidence ( 13 terms )
Molecular Function
CV term Evidence References
RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in negative regulation of transcription inferred from direct assay (Sanicola et al., 1995) RNA polymerase II distal enhancer sequence-specific DNA binding inferred from direct assay (Sanicola et al., 1995)
Biological Process
CV term Evidence References
anterior commissure morphogenesis inferred from mutant phenotype (Bossing and Brand, 2006) axon guidance inferred from mutant phenotype (Bossing and Brand, 2006) genital disc anterior/posterior pattern formation inferred from expression pattern (Freeland and Kuhn, 1996) genital disc development inferred from mutant phenotype (Chen et al., 2005) gonad development inferred from mutant phenotype (Weyers et al., 2011) negative regulation of gene expression inferred from mutant phenotype (Seibert and Urbach, 2010) negative regulation of transcription from RNA polymerase II promoter inferred from mutant phenotype (Sanicola et al., 1995) neuroblast fate determination inferred from genetic interaction with inv, gsb AND inferred from genetic interaction with inv, gsb, ptc (Bhat and Schedl, 1997) spiracle morphogenesis, open tracheal system inferred from mutant phenotype (Merabet et al., 2005) ventral midline development inferred from mutant phenotype (Bossing and Brand, 2006)
Cellular Component
CV term Evidence References
nucleus inferred from direct assay (DiNardo et al., 1985)
Terms Based on Predictions or Assertions ( 18 terms )
Molecular Function
CV term Evidence References
sequence-specific DNA binding transcription factor activity inferred from sequence or structural similarity with UniProtKB:Q05925 (FlyBase, 1992-)
Biological Process
CV term Evidence References
analia development traceable author statement (Sanchez and Guerrero, 2001) anterior/posterior lineage restriction, imaginal disc traceable author statement (Milan and Cohen, 2000, McNeill, 2000) anterior/posterior pattern specification, imaginal disc traceable author statement (Marsh and Theisen, 1999) anterior head segmentation traceable author statement (Peel, 2004) central nervous system development non-traceable author statement (Swiss-Prot Project Members, 1986.7.21) compartment pattern specification non-traceable author statement (Calleja et al., 2002) traceable author statement (Milan and Cohen, 2000) genital disc anterior/posterior pattern formation traceable author statement (Sanchez and Guerrero, 2001) imaginal disc-derived female genitalia development traceable author statement (Sanchez and Guerrero, 2001) imaginal disc-derived male genitalia development traceable author statement (Sanchez and Guerrero, 2001) imaginal disc pattern formation traceable author statement (Potter and Xu, 2001) negative regulation of transcription, DNA-dependent non-traceable author statement (Swiss-Prot Project Members, 1986.7.21, Courey and Jia, 2001) posterior compartment specification traceable author statement (McNeill, 2000) posterior head segmentation traceable author statement (Peel, 2004) segment polarity determination non-traceable author statement (Swiss-Prot Project Members, 1986.7.21, Canon and Banerjee, 2000, Sanson, 2001, Davis and Patel, 2003) traceable author statement (Davies and Patel, 1999, Peel, 2004) trunk segmentation traceable author statement (Peel, 2004) wing disc anterior/posterior pattern formation traceable author statement (Milan and Cohen, 2000, McNeill, 2000)
Cellular Component
CV term Evidence References
nucleus non-traceable author statement (Swiss-Prot Project Members, 1986.7.21)
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 en allele Gene References en1 ci (Locke and Hanna, 1996) ph-p (Maschat et al., 1998) en4 ci (Locke and Hanna, 1996) eRF1 (Chao et al., 2003) ph-d (Maschat et al., 1998) ph-p (Maschat et al., 1998) en7 eRF1 (Chao et al., 2003) en9.6 inv (Tabata et al., 1995) en54 ph-d (Maschat et al., 1998) ph-p (Maschat et al., 1998) en58 inv (Whiteley and Kassis, 1997) en59 cg (Campbell and Tomlinson, 2000) ci (Locke and Hanna, 1996) en96 dpp (Sanicola et al., 1995) enC ph-p (Maschat et al., 1998) enE inv (Bhat and Schedl, 1997, Takashima and Murakami, 2001) enEnci ci (Locke and Hanna, 1995, Locke and Hanna, 1996) enunspecified wg (Gritzan et al., 1999) enXho25 ph-p (Maschat et al., 1998) unspecified cg (House, 1953, House, 1953, House, 1961, Mukherjee, 1965, Datta and Mukherjee, 1971, Svendsen et al., 2000) ci (House, 1953, House, 1953, House, 1961, Mukherjee, 1965, Datta and Mukherjee, 1971, Dahmann and Basler, 2000) fu (Lawrence and Struhl, 1982) hh (Maschat et al., 1998) nub (Cifuentes and Garcia-Bellido, 1997) rho (Sturtevant and Bier, 1995) smo (Dahmann and Basler, 2000) sx (House, 1953, House, 1953, House, 1961, Mukherjee, 1965, Datta and Mukherjee, 1971)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 62028 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0000577 DroID - A comprehensive database of gene and protein interactions. • FBgn0000577 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 36240
Orthologs
Genome-wide drosophilid orthologs	Dana\GF12393 Dere\GG22644 Dgri\GH20500 Dmoj\GI21047 Dper\GL10677 Dpse\en Dsec\en Dsim\en Dvir\en Dwil\GK22225 Dyak\GE13516
Curated drosophilid orthologs	Dacu\en   Dant\en   Dark\en   Darw\en   Dawd\en   Dcai\en   Dddt\en   Dddu\en   Dhyd\en (Spencer, 1942) Dmir\en   Dngo\en   Dore\en (Dermitzakis et al., 2001) Dpic\en   Dpse\en (Schaeffer and Anderson, 1997) Dsec\en (Dermitzakis et al., 2001) Dsig\en   Dsii\en   Dsim\en (Dermitzakis et al., 2001) Dvir\en   Dyak\en (Dermitzakis et al., 2001)
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL014635-PA Aedes aegypti (mosquito) AGAP008023-PA Anopheles gambiae (mosquito) BGIBMGA009797-PA Bombyx mori (silk worm) JA12584 Caenorhabditis japonica (nematode) CPIJ010220 Culex pipens (mosquito) 290629 Daphnia pulex (daphnia) 290638 Daphnia pulex (daphnia) ENSDARP00000012379 Danio rerio (zebra fish) ENSGACP00000019729 Gymnopilus aculeatus (mushroom) ENSMODP00000006129 Monodelphis domestica (gray opossum) ENSMMUP00000010208 Macaca mulatta (rhesus monkey) XP_001607672.1 Nasonia vitripennis (wasp) ENSOANP00000014598 Ornithorynchus anatinus (platypus) ENSORLP00000025789 Oryzias latipes (japanese killifish) LOC_Os10g23090.1 Oryza sativa (asian rice) LOC_Os10g26500.1 Oryza sativa (asian rice) PHUM006221-PA Pediculus humanus (human body louse) ENSPTRP00000021643 Pan troglodytes (chimpanzee) XP_001189948.1 Strongylocentrotus purpuratus (purple sea urchin) XP_972888.1 Tribolium castaneum (red flour beetle) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • NV10873 LH20848 LH20842 ISCW022797 FBgn0231183 FBgn0230114 FBgn0224210 FBgn0224059 FBgn0013111 FBgn0182565 FBgn0044189 FBgn0176176 FBgn0044210 JGI_V11_290629 JGI_V11_290630 JGI_V11_290638 FBgn0074717 FBgn0020475 FBgn0149024 FBgn0148287 FBgn0143779 FBgn0001269 FBgn0130996 FBgn0114815 FBgn0089596 FBgn0089430 CPIJ010220 BGIBMGA009798 BGIBMGA009644 ACYPI008905 ACYPI002874 GB15566 HMEN_ANOGA AAEL014635 PB19748 PB19750 PHUM030340 GLEAN_08952
Stocks & Reagents
Stocks Listed in FlyBase ( 38 )
Bloomington	410 enspt 1287 enApa/SM6#16 3516 P{hs-en3}3; ry* 265 cn1 en1/SM5 1817 cn1 en4/SM5 1820 cn1 en7/SM5 256 chl1 en1/SM5
Kyoto	101487 enApa / SM6#16 105822 cn1 en1/SM5 106520 cn1 en4/SM5 106523 cn1 en7/SM5
VDRC	v105678 P{KK107874}VIE-260B
	More stocks available...
Genomic Clones ( 1 )
	BACR09A11 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 99 )
	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	LD16125
Other clones	DPiM_LD16125
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	AT28921 RH68203
Other clones	12688 58395 71538 75331 81253 98286 214313.22 214319.23 CK01.53A.F2 CK01.53D.G1 CK01.65B.C2 CK01.105D.D3 CK01.123D.E9 EK042853 EK043425 EK053141 EK088414 EK181347 EK199433 EK202607 EK205355 EP09447 EP10964 IP07553 RP004019366 RP004047072 RP004053711 RP004058266 RP004065012 RP004074872 RP004078522 RP004108506 RP004113808 RP004120774 RP004130271 RP004146220 RP004147408 RP004162403 RP004175219 RP004177181 RP004178655 RP004197621 RP004217344 RP004221463 RP004221736 RP004230033 RP004231554 RP004232066 RP004232865 RP004233757 RP004238772 RP004241972 RP004244249 RP004246651 RP004247956 RP004248780 RP004253399 RP004254666 RP004257224 RP004258021 RP004260951 RP004261034 RP004261971 RP004267914 RP004299798 RP004311286 RP004321531 RP004325565 RP004326348 RP004326449 RP004344637 RP004345598 RP004359595 RP004368509 RP004374585 RP004387262 RP004394499 RP004416926 RP004437506 RP004439882 RP004458823 RP004460461 RP004476362 RP004480041 RP004491923 RP004496564 RP004506043 RP004506480 RP004517250 RP004527075 RP004533569 RP004538108 RP004544228 RP004545410 RP004550905
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0000577 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 36240
Antibody Information
	monoclonal (Patel et al., 1989) monoclonal antibody (Larsen et al., 2003) polyclonal (DiNardo et al., 1985) polyclonal antibody (Chanas et al., 2004)
Other Information
Discoverer
Etymology
	The gene is named "engrailed", a heraldic term from the middle-age french "engraile", literally "dented by hail", after the mutant phenotype of a notch in the scutellum. (Gibert, 2002)
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of
Additional comments	The highly divergent mosquito engrailed protein Agam\en is able to substitute for en protein during all stages of development. Agam\en expression by enhancer-trapping is precise enough to restore viability to en mutants. (Whiteley and Kassis, 1997) enspt fails to complement en1. (Chase and Baker, 1995)
Other Comments
	Ectopic expression of en prior to neuronal specification can misroute axons, whereas en has no major effect when ectopically expressed in neurons only. Therefore, the action of en on axonal pathfinding occurs at early stages, before neuronal cell fate is established. (Joly et al., 2007) A 139bp minimal pairing sensitive element (PSE) from en regulatory DNA has been defined. This element mediates Pairing Sensitive Silencing (PSS). The PSE includes 8 protein binding sites at least 5 of which are important for pairing sensitive silencing. One of these is for pho protein, and another has the GAGAG sequence known to be bound by Trl and psq proteins. (Americo et al., 2002) fz and fz2 are required during embryogenesis to maintain epidermal en and wg expression. (Bhanot et al., 1999) hh is required for activation of en during regeneration of fragmented imaginal discs. (Gibson and Schubiger, 1999) wg mutant embryos have residual mirror-symmetric pattern due to an en-dependent signal specifying anterior denticle fates. The en-dependent signal acts unidirectionally and wg activity imposes the asymmetry. (Gritzan et al., 1999) Three EMS induced en alleles were identified in a screen for mutations affecting commissure formation in the CNS of the embryo. (Hummel et al., 1999) In the developing abdomen the state of en, whether off or on, determines whether a cell is of Anterior or Posterior type. en acts in the Anterior compartment, where it is induced by hh gene product, in a narrow band of cells, which has crossed the Anterior/Posterior compartment boundary. en causes these cells to form a special type of cuticle. (Lawrence et al., 1999) Cell affinities in the adult abdomen depend on en: cells of the P compartment have affinities that are distinct from A cells. (Lawrence et al., 1999) When en is removed from Posterior cells, they transform into the Anterior cells of the same parasegment, rather than Anterior cells of the same segment. (Lawrence et al., 1999) Transport or stability of wg gene product is reduced within en expressing cells. (Sanson et al., 1999) pho encodes a sequence-specific DNA binding protein that interacts with PRE in the en gene. (Brown et al., 1998) In solution the en homeodomain binds to a DNA molecule containing a single TAAT sequence in an analogous manner to that observed in the X-ray co-crystal structure. en also binds to a more complex site, having overlapping TAAT sequences, in more than one orientation. (Draganescu and Tullius, 1998) eg is necessary for the maintenance of en and zfh2 expression in the serotonin neurons. (Lundell and Hirsh, 1998) Interactions between en and polyhomeotic are required to maintain the anterior-posterior boundary and posterior cell fate in the wing. (Maschat et al., 1998) Segment polarity gene expression is necessary for the survival of specific rows of epidermal cells. (Pazdera et al., 1998) en promotes the development of the dorsolateral fat body, midgut visceral mesoderm and somatic gonads while it suppresses development of somatic muscles, heart and ventral fat body. There is a balance between fat body and somatic gonadal precursor (SGP) development with tin, wg and en driving cells in the primary clusters towards SGP development and srp driving them towards fat body development. (Riechmann et al., 1998) Multiple domains contribute to en protein repression activity. The eh1 region interacts with the gro repressor. (Tolkunova et al., 1998) CrebA, en double mutant phenotype confirms that CrebA is not involved in segment polarity. (Andrew et al., 1997) Either en or inv is required to uncouple the ptc-gsb regulatory circuit in row 5 neuroblasts during neurogenesis. (Bhat and Schedl, 1997) Genes known to be expressed in compartment-specific manner in discs are expressed in analogous patterns in each primordia. (Chen and Baker, 1997) Adjacent and conserved ftz and cofactor binding sites within the en intron enhancer are necessary and sufficient for transcriptional activation. The cofactor sites can be specifically bound by ftz-f1, and the ftz homeodomain and ftz-f1 bind cooperatively in vitro. (Florence et al., 1997) Cross-regulatory relationships among hh, wg and en, as well as their initial mode of activation, in the anterior head are significantly different from those in the trunk. (Gallitano-Mendel and Finkelstein, 1997) pros does not modulate en regulatory region binding to DNA. (Hassan et al., 1997) The expression of wg and en in the adult antenna is controlled by age-dependent mechanisms. (Rogina and Helfand, 1997) βTub60D is directly repressed by en protein. The first intron of the βTub60D locus contains several en protein binding sites. (Serrano et al., 1997) Pc, Psc and polyhomeotic proteins are associated with identical regulatory elements of en in tissue culture cells and differentially distributed on regulatory sequences of inv. (Strutt and Paro, 1997) en is one of a class of genes with TATA-less promoters that have the conserved DPE sequence. (Burke and Kadonaga, 1996) Amino acids in the N-terminal arm of the homeodomain, as well as at position 54 of the homeodomain, control the DNA binding specificity of the homeodomain. (Damante et al., 1996) Cells in anterior compartments lacking ci express hh and adopt a posterior fate without expressing en. (Dominguez et al., 1996) ara-caup are expressed at patches on the wing, located one at each side of the DV compartment border. The posterior border of the patches is defined by repression by en. (Gomez-Skarmeta and Modolell, 1996) Despite the absence of a syncytium in C.floridanum embryos monoclonal antibodies to en, Ubx and abd-A demonstrate their cognate proteins are expressed in a conserved pattern in the post-gastrulation stages of development. The expression of the eve cognate protein is not completely conserved and lacks a pair rule phase to its expression. (Grbic et al., 1996) The varied phenotypes of en alleles can be explained by the differential effects these mutants have on the combination of en and inv activities. (Gustavson et al., 1996) en and inv share a regulatory region and encode redundant functions. (Gustavson et al., 1996) Interfaces between posterior (en on) and anterior (en off) cells are required for pattern formation. wg could play the role of the morphogen, at least within part of the segmental pattern. (Lawrence et al., 1996) Four segment polarity genes, hh, wg, gsb and en all function in concert to determine the formation and specifications of three hh-dependent eg-neuroblasts (6-4, 7-3 and 2-4). (Matsuzaki and Saigo, 1996) Elevated en expression does not disrupt eye morphogenesis. (Strutt and Mlodzik, 1996) en is not required for hh activation or maintenance in the eye disc. Elevated en levels repress dpp, ptc and ci expression. (Strutt and Mlodzik, 1996) en is post-translationally modified by casein kinase II (CK-II). The targets for CK-II phosphorylation in vitro are serine residues 394, 397, 401 and 402, and phosphorylation by CK-II stimulates DNA binding. (Bourbon et al., 1995) Maintenance but not initiation of en gene expression in the embryo requires trx, which is also required to maintain stable long-term expression of the homeotic genes throughout development. trx is required for normal en expression in the wing imaginal disc. trx-dependent loss of en expression in the dorsal fat body is correlated with female sterility. (Breen et al., 1995) gro and hh regulate en expression in the anterior compartment of the wing. (de Celis and Ruiz-Gomez, 1995) Early ph-p gene expression is under the control of bcd and en as activators and of osk as an inhibitor. (Fauvarque et al., 1995) Early eve expression is required for the activation of both even and odd numbered en stripes and late eve stripes strengthen the expression of odd numbered en stripes. (Fujioka et al., 1995) en has a dual role in wing development: a general role in patterning the appendage, achieved through the activation of secreted proteins like hh and dpp, and a more specific one, determining posterior identity, in which the inv gene may be implicated. (Guillen et al., 1995) The organisation of the tail region of the embryo is documented from studies of cuticular markers enabling a more direct comparison between homologous structures on the embryo and larval cuticle. (Kuhn et al., 1995) en+ is a dose-dependent modifier of the ci locus. Lack of pairing at the ci locus can facilitate the en--dependent expression of the ci phenotypes. (Locke and Hanna, 1995) en is involved in regulation of serotonin cell development. (Lundell and Hirsh, 1995) odd and nkd are required to restrict en expression. odd represses expression of ftz, an activator of en. nkd prevents activation of en by ftz without affecting ftz expression. (Mullen and DiNardo, 1995) The transition from inter-dependent wg and en expression to wg autoregulation may involve several segment polarity genes. (Noordermeer et al., 1995) en is a candidate oc target gene in ocelli formation. (Royet and Finkelstein, 1995) Msr-110 is a direct target for regulation by en. Normal expression of Msr-110 is dependent upon en function. (Saenz-Robles et al., 1995) Clonal analysis supports the view that dpp is a direct target of repression by en, and that en defines the posterior extent of the dpp stripe in the wing imaginal disc. The en-hh-ptc regulatory loop that is responsible for segmental expression of wg in the embryo is reused in imaginal disks to create a stripe of dpp expression along the A/P compartment boundary. (Sanicola et al., 1995) ph-d and ph-p activation in germ band elongated embryos could be mediated by en binding to en-binding sites upstream of each of the ph-d and ph-p transcription units. (Serrano et al., 1995) inv can perform a subset of functions attributed to en including the ability to confer posterior compartment identity, but does not appear to organize the anterior-posterior compartment boundary. (Simmonds and Bell, 1995) The establishment of the anterior-posterior organiser and control of compartment identity are genetically distinguishable and inv may perform a discrete subset of functions previously ascribed to en. (Simmonds et al., 1995) en both directs the posterior compartment pathway and creates the compartment border in wing development. (Tabata et al., 1995) Maintenance of restricted wg transcription during late gastrulation requires en-independent wg autoregulatory activity. (Yoffe et al., 1995) en governs growth and patterning in both anterior and posterior wing compartments by controlling the expression of the hh and dpp products as well as the response of the cells to them. en activity programs wing cells to express hh whereas the absence of en activity programs them to respond to hh by expressing dpp. Consequently, posterior cells secrete hh product and induce a stripe of neighboring anterior cells across the compartment boundary to secrete dpp. (Zecca et al., 1995) opa activity is essential for the appropriate level and timing of en and wg expression in all parasegments, but does not determine their restricted spatial domains of expression. (Benedyk et al., 1994) Clones lacking en function can respect a boundary of lineage restriction. hh mediates the interaction between anterior dpp-expressing cells and posterior en-expressing cells. (Hidalgo, 1994) en is required for patterning of the posterior wing margin, specifically to inhibit the formation of stout and socketed bristles (characteristic of the anterior wing margin) posterior to wing vein 3. en participates, presumably indirectly, in cell-cell interactions. (Hidalgo, 1994) Three pairing sensitive (PS) sites map to a 1.6kb fragment of en. PS sites of en can suppress expression of a linked marker gene (w) carried in P-elements, dependent on the presence of two copies of the P-element in proximity in the genome. (Kassis, 1994) en pairing sensitive (PS) sites apparently normally act to promote interactions between distantly located en regulatory sites and the en promoter. (Kassis, 1994) wg product made in the mesoderm can sustain en expression in the ectoderm. (Lawrence et al., 1994) en-mediated repression of Ubx is necessary for the parasegment 6 identity. (Mann, 1994) wg acts through dsh and arm to affect the expression of en and cuticle differentiation. (Noordermeer et al., 1994) Comparisons of early development to that in other insects have revealed conservation of some aspects of development, as well as differences that may explain variations in early patterning events. (Patel, 1994) The exd protein modulates the DNA binding activity of en to a different site. In direction of increasing cytology: inv+ en- (van Dijk and Murre, 1994) Maintenance of en expression does not always require wg activity. (Bejsovec and Wieschaus, 1993) Wild type activity of five segment polarity genes, wg, ptc, en, nkd and hh, can account for most of the ventral pattern elements in the embryo. wg is required for naked cuticle. wg generates the diversity of cell types within the segment but each specific cell identity depends on the activity of ptc, en, nkd and hh. (Bejsovec and Wieschaus, 1993) wg functions as an extracellular signal maintaining en expression. (Cumberledge and Krasnow, 1993) en is a specific repressor of activated transcription, and may act via a different mechanism than eve, perhaps by interfering with interactions between transcriptional activators and the general transcription machinery. A minimum repression domain of 55 residues, rich in alanine, can function when fused to a heterologous DNA binding domain. Unlike the repression domains of eve and Kr, the en repression domain is moderately charged. (Han and Manley, 1993) A temporal asymmetry in the regulatory interactions between wg, en and gsb explains the difference in phenotypes between the mutants. (Li and Noll, 1993) The positioning of en stripes in the embryo is largely determined by the actions of negative regulators: run is required to limit the domains of en-expression in odd-numbered parasegments, while odd is required to limit the domains of en-expression in even-numbered parasegments. Activation of en at the anterior margins of the parasegments requires repression of run and odd by eve. (Manoukian and Krause, 1993) The BRE region of Ubx includes binding sites for hb, ftz, tll, en and twi. The binding of their products and the interplay between them is responsible for generating the expression pattern directed by the BRE. (Qian et al., 1993) Overexpression of wg protein does not prevent parasegmental, restricted en expression, therefore wg is not acting as a local instructive signal or a morphogen. (Sampedro et al., 1993) Choice of cell fate made by en expressing cells in embryonic parasegments is mediated by wg, in a function distinct from its early role in maintaining en expression. en expressing cells respond differently to wg at different stages of development:early wg stabilizes the subdivision of the body axis by maintaining en expression, whereas later input generates cell-type diversity. (Dougan and DiNardo, 1992) Experiments designed to identify cis-acting regulatory elements of en revealed a tendency for fragments of en DNA to target insertions to genes that are expressed in stripes. (Kassis et al., 1992) en alleles fall into four classes, all recessive. Class 1: en1 (See en1 allele record for full description). Viable hemizygous and homozygous; fertile. Longitudinal cleft extends from rear border of scutellum forward, may be reduced to median nick or posterior flattening of scutellum. Wings larger, broader and thin textured with spatulate end; venation and distribution of sensilla abnormal in posterior wing compartment. Variable duplication of anterior triple row bristles on posterior margin; alula reduced, with costal-like bristles. Clones of en1 cells of posterior compartment origin fail to respect anterior-posterior compartment border in wing disc as do mwh1 clones in wing discs of en1 homozygotes. en1 abnormalities are associated with posterior compartment structures only, except for scutellar cleft. Not suppressed by su(Hw)1. Class 2: Lethal alleles with normal cytology. Embryonic lethal. Anterior margin of each segment defective. Pair rule defects in naked cuticle of T1, T3, A2, A4, A6, A8 result in pair-wise fusion of adjacent segments. Autonomous effects in adult cuticular clones observed in posterior compartment of proboscis, thorax, abdomen and genitalia. The en1/"enlethal" heterozygote characterized by wing abnormalities only; disruption of anterior crossvein, gap in vein IV and occasional socketted bristles on posterior margin. Class 3: Deficiencies and lethal alleles with inversion or translocation breakpoints. Embryonic lethal. Embryonic segment defects slight and variable. Alleles of this class in heterozygous combination witn en1 produce adults more extreme than en1. For example, in en1/en2, legs are truncated, the tarsi reduced to densely bristled stumps; wings are greatly enlarged and spatulate with greater disruption of veins IV and V; higher penetrance of socketed bristles along the posterior margin. Extreme scutellar cleft. Class 4: Non-lethal alleles with breakpoints. Hemizygous viable, embryos normal. Heterozygous with other allele classes, variable gaps in wing veins IV and V. Variable reductions or deletions in male and female genitalia. Scutellum may also be affected. (Lindsley and Zimm, 1992) en is activated by ftz and repressed by odd, and low levels of eve cause ftz-dependent activation of en through repression of odd. (Manoukian and Krause, 1992) Embryos mutant for two or more Pc-group genes (Pc, Scm, Pcl, Psc, Asx, E(Pc), E(z), ph-d, pho and esc) show strong ectopic en expression, but only weak derepression occurs if embryo is mutant at only one of the Pc group genes. This effect is independent of the function of en itself, and wg. Expression of en in esc mutant embryos is almost normal, suggesting that esc may function in a pathway different from the other genes in the group. (Moazed and O'Farrell, 1992) Ectopic uniform wg expression results in expansion of the en expression domain in a posterior direction: original parasegment borders remain unchanged. (Noordermeer et al., 1992) The 88kD ttk protein isoform binds a site upstream of the en promoter. (Read and Manley, 1992) wg signalling operates by inactivating the sgg repression of en autoregulation. (Siegfried et al., 1992) Expression pattern of hh coincides with that of en in the epidermis. Though initially independent of en, hh expression later becomes en-dependent. In the absence of ptc function, wg expression, which is normally en-dependent, no longer requires en. (Tabata et al., 1992) Clonal analysis shows that no clones straddle the anterior edge of en stripes in the embryo. However, posterior cells of each stripe lose en expression, producing mixed clones. (Vincent and O'Farrell, 1992) wg and en function in patterning the larval epidermis. (Bejsovec and Martinez Arias, 1991) In embryos homozygous for sm, en expression fades towards the end of the extended germband phase suggesting that sm is important in maintaining en after its initial activation. (Blader et al., 1991) en expression pattern in the embryonic head strengthens the hypothesis that the clypeolabrum evolved from the fusion of paired labral appendages. (Diederich et al., 1991) en expression becomes independent of wg extracellular influence and relies on positive autoregulation. Autoregulation is transient so does not supply a mechanism for determination of the en-cell fate. (Heemskerk et al., 1991) en cannot completely rescue the ptc phenotype when in double mutant combinations. (Hidalgo, 1991) In transfection assays en is an active transcriptional repressor. Active repression is distinct from the effects of passive homeodomain-containing proteins which repress when competing with activators for binding sites and activate when competing with en. Active repression activity maps outside the en homeodomain and this activity can be transferred to a heterologous DNA binding domain. (Jaynes and O'Farrell, 1991) When a fragment of en DNA extending from -2.4kb to +0.8kb is included in the CaSpeR vector the eye colour of transformants often behaves anomalously, half of the homozygotes have a lighter eye colour than the heterozygotes and a quarter have the same eye colour as the heterozygotes. This occurred at many different insertion sites whether the upstream en fragment is directly adjacent to w+mC or when it is 1kb away. The suppression of eye colour is dependent on the proximity of the two copies of the transposon, either in cis or trans. The protein that mediates this phenomenon is probably not a homeodomain-containing protein as deletion of the homeodomain binding sites results in suppression of w expression. (Kassis et al., 1991) Mutations in zygotic polarity gene en do not interact with RpII140wimp. (Parkhurst and Ish-Horowicz, 1991) Appropriate en activity is required for proper restriction of the dpp expression domain as well as being required for maintenance of the posterior compartment fate. (Raftery et al., 1991) en represses ciD expression in the posterior compartments of embryos and imaginal discs. (Eaton and Kornberg, 1990) en mutants exhibit variable pair rule fusions and segment polarity reversals. (Gaul and Jaeckle, 1990) ptc is negatively regulated by en in the early extended germ band. (Hidalgo and Ingham, 1990) en intron 1 works as a stripe specific enhancer. (Kassis, 1990) The crystal structure of a complex containing the en homeodomain and a duplex DNA site has been determined. (Kissinger et al., 1990) A modification and reduction in en and Dfd protein distribution is seen in mutant cad embryos. (Mlodzik et al., 1990) sim gene product is required for the normal expression of en. (Nambu et al., 1990) en protein represses transcriptional activation by ftz protein by competition for binding to homeodomain binding sites in vitro. (Ohkuma et al., 1990) The en1/"enlethal" heterozygote characterized by wing abnormalities only; in some combinations complementation is complete or nearly so. (Condie and Brower, 1989) A transient expression assay has been employed to investigate the potential of homeobox genes to function as transcriptional activators. (Han et al., 1989) A monoclonal antibody (MAb4D9) reacts specifically to en proteins in a variety of species. An ancestral function of en may have been in neurogenesis and its other functions in arthropods may represent a more recent addition. (Patel et al., 1989) The expression of en protein in Drosophila, grasshopper and crayfish has been compared. (Patel et al., 1989) Genetic analysis demonstrates that en is dispensable for efficient homeotic gene expression in the visceral mesoderm. (Tremml and Bienz, 1989) Scr and en are derepressed in the absence of Pc and the bithorax complex function. (Busturia and Morata, 1988) Striped en expression during post-blastoderm development is controlled by a cis-regulatory programme distinct from that controlling the establishment of expression at the cellular blastoderm stage. (DiNardo et al., 1988) en protein isolated from cultured cells and embryos is post-translationally modified. (Gay et al., 1988) Heat shock induced en gene expression causes pattern defects similar to those in embryos lacking en gene product. The sensitivity to heat shock is only during the cellular blastoderm and early gastrulation periods when the en protein localises into segmentally reiterated stripes and represents only a small portion of the normal period of en gene expression. (Poole and Kornberg, 1988) en is expressed in the posterior (but not the anterior) compartments of the embryo and larva. (Kornberg et al., 1985) en mutant genital discs show reductions or deletions. (Epper and Sanchez, 1983) "enlethal" alleles show no maternal effect. (Lawrence et al., 1983) At 29oC, duplication of anterior compartment bristles in mirror-image symmetry in the posterior compartment of second antennal segment occurs. (Morata et al., 1983) "enlethal" clones are without effect in anterior compartments and in eye-antennal region. (Kornberg, 1981, Kornberg, 1981, Lawrence and Struhl, 1982)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_078976 NM_165841 RefSeq (Proteins) • NP_523700 NP_725059 DDBJ / EMBL / GenBank • AA441254 AAA65478 AAF58639 AAF58639.1 AAL39593 AAM68711 AAM68711.1 AW942397 AY069448 AY184070 AY184071 AY184072 AY184073 AY184074 AY184075 AY184076 AY184077 AY184078 AY184079 AY184080 AY184081 CAA25906 K03055 K03056 M10017 M29285 X01765 Entrez Gene - A searchable database of RefSeq genes. • 36240 UniProtKB/Swiss-Prot • P02836
Other Crossreferences
	EPD - Eukarytoic Promoter Database, an annotated collection of POL II promoters • 24009 InterPro domains - A database of protein families, domains, and functional sites • Homeobox, conserved site (IPR017970) Helix-turn-helix motif, lambda-like repressor (IPR000047) Homeobox engrailed (IPR000747) Homeobox (IPR001356) Homeodomain-like (IPR009057) Homeodomain-related (IPR012287) Homeobox engrailed, C-terminal (IPR019549) Homeobox engrailed-type, conserved site (IPR019737) Homeobox, eukaryotic (IPR020479) PDB - Protein Data Bank. An information portal to biological macromolecular structures • 1ENH 1HDD 2HDD 3HDD
Linkouts
	BioGRID - A database of protein and genetic interactions • 62028 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0000577 DroID - A comprehensive database of gene and protein interactions. • FBgn0000577 DRSC - Results from RNAi screens. • FBgn0000577 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0000577 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG9015-RB FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0000577 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 36240 InParanoid A subset of ortholog calls from InParanoid. • AAEL014635-PA Aedes aegypti (mosquito) AGAP008023-PA Anopheles gambiae (mosquito) BGIBMGA009797-PA Bombyx mori (silk worm) JA12584 Caenorhabditis japonica (nematode) CPIJ010220 Culex pipens (mosquito) 290629 Daphnia pulex (daphnia) 290638 Daphnia pulex (daphnia) ENSDARP00000012379 Danio rerio (zebra fish) ENSGACP00000019729 Gymnopilus aculeatus (mushroom) ENSMODP00000006129 Monodelphis domestica (gray opossum) ENSMMUP00000010208 Macaca mulatta (rhesus monkey) XP_001607672.1 Nasonia vitripennis (wasp) ENSOANP00000014598 Ornithorynchus anatinus (platypus) ENSORLP00000025789 Oryzias latipes (japanese killifish) LOC_Os10g23090.1 Oryza sativa (asian rice) LOC_Os10g26500.1 Oryza sativa (asian rice) PHUM006221-PA Pediculus humanus (human body louse) ENSPTRP00000021643 Pan troglodytes (chimpanzee) XP_001189948.1 Strongylocentrotus purpuratus (purple sea urchin) XP_972888.1 Tribolium castaneum (red flour beetle) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /segment/engrail1.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 36240 modMine - Data generated by the modENCODE project. • FBgn0000577 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • NV10873 LH20848 LH20842 ISCW022797 FBgn0231183 FBgn0230114 FBgn0224210 FBgn0224059 FBgn0013111 FBgn0182565 FBgn0044189 FBgn0176176 FBgn0044210 JGI_V11_290629 JGI_V11_290630 JGI_V11_290638 FBgn0074717 FBgn0020475 FBgn0149024 FBgn0148287 FBgn0143779 FBgn0001269 FBgn0130996 FBgn0114815 FBgn0089596 FBgn0089430 CPIJ010220 BGIBMGA009798 BGIBMGA009644 ACYPI008905 ACYPI002874 GB15566 HMEN_ANOGA AAEL014635 PB19748 PB19750 PHUM030340 GLEAN_08952 REDfly - A database of transcriptional regulatory elements. • FBgn0000577
Synonyms & Secondary IDs ( 22 )
Reported As
Symbol Synonym	153867_at (Kristensen et al., 2005) Apa   CG9015 (Perkins et al., 2009.3.31, Keleman et al., 2009.8.5, Ni et al., 2010.12.1) en (Casso et al., 2008, Sanders et al., 2008, Kategaya et al., 2009, Peel et al., 2005, Urbach et al., 2006, Yasunaga et al., 2006, Chanas and Maschat, 2005, Srivastava et al., 2002, Gorfinkiel et al., 2005, Wang and Gergen, 2007, Jaeger and Reinitz, 2006, Muro et al., 2006, McEwen and Peifer, 2005, Mosimann et al., 2006, Croker et al., 2006, Buescher et al., 2006, Sanchez-Elsner et al., 2006, Freeland and Kuhn, 1996, Molnar and de Celis, 2006, Wheeler et al., 2006, Kent et al., 2006, Bossing and Brand, 2006, Yao et al., 2006, Bartolome and Charlesworth, 2006, Kannan et al., 2010, Gallet et al., 2006, Ntini E and Wimmer, 2007, Sandmann et al., 2007, Sprecher et al., 2006, Scholtz, 2006, Beckett and Baylies, 2006, Bowler et al., 2006, Aerts et al., 2007, Venken et al., 2009, Kremer et al., 2010, Park et al., 2008, Negre et al., 2006, Liu et al., 2005, Deshpande and Schedl, 2005, Urbach, 2007, Molnar et al., 2011, Peel et al., 2006, Crickmore and Mann, 2007, Wendler et al., 2006, Tarone et al., 2008, Durant and Kassis, 2008, Bejarano et al., 2005, Maeda et al., 2007, Brodsky et al., 2004, Torroja et al., 2005, Shen et al., 2008, Von Ohlen et al., 2007, Jamieson et al., 2006, Wang et al., 2007, Ringrose and Paro, 2007, Liu et al., 2007, Goldstein et al., 2005, Vanderzwan-Butler et al., 2007, Joly et al., 2007, Bolivar et al., 2006, Lee et al., 2007, Sgaier et al., 2007, Chan et al., 2007, Chan et al., 2008, Nekrasov et al., 2007, Comet et al., 2006, Gallet et al., 2008, Ruel et al., 2007, Coiffier et al., 2008, Haecker et al., 2008, Umemori et al., 2007, Gebelein and Mann, 2007, Claret et al., 2007, Mulinari et al., 2008, Kim et al., 2008, Hendrix et al., 2008, Siera and Cline, 2008, Dominguez-Gimenez et al., 2007, Schuettengruber et al., 2009, Garaulet et al., 2008, Larsen et al., 2008, Gaziova and Bhat, 2009, Wheeler et al., 2009, Bejarano and Milán, 2009, Stagg et al., 2011, Fang et al., 2009, Grimm et al., 2009, Bejarano et al., 2007, Larsen et al., 2009, Kwon et al., 2009, Bosveld et al., 2008, Mosimann et al., 2009, Gambetta et al., 2009, Gambetta et al., 2009, Zhai et al., 2009, Schwartz et al., 2006, Younossi-Hartenstein et al., 2006, Glise et al., 2005, True et al., 2001, Meier et al., 2006, Hou et al., 2009, Seibert et al., 2009, Mulinari and Häcker, 2009, Kumar et al., 2009, Guo and Wang, 2009, Cheng et al., 2010, Beckervordersandforth et al., 2008, Grieder et al., 2009, Landsberg et al., 2009, Venken et al., 2009, Bischoff et al., 2009, Colomb et al., 2008, Grad et al., 2004, Klein et al., 2010, Wang et al., 2010, Rousset et al., 2010, Braid et al., 2010, Hartmann et al., 2010, Blagburn, 2008, Gettings et al., 2010, Monier et al., 2010, Xu et al., 2008, González et al., 2008, Gurunathan et al., 2004, Wang et al., 2010, Jones et al., 2010, Ding et al., 2010, Walrad et al., 2010, Lundell et al., 1996, Menéndez et al., 2010, Cunningham et al., 2010, Schwartz et al., 2010, Yassin et al., 2010, Jung et al., 2010, Lee et al., 1999, Walrad et al., 2011, Benitez et al., 2010, Beltran et al., 2007, Subramanian and Gadgil, 2010, Klebes et al., 2002, Terriente-Félix et al., 2011, Fiedler et al., 2011, Watson et al., 2011, Hogan et al., 2011, Chou et al., 2010, Seibert and Urbach, 2010, Weyers et al., 2011, Sprecher and Hirth, 2006, Jia et al., 2010, Kazemian et al., 2010, Tchuraev and Galimzyanov, 2009) En (Mosimann et al., 2009, Kategaya et al., 2009, Goodman et al., 2006, Wehn and Campbell, 2006, Zhou et al., 2006, Waldrop et al., 2006, Noro and Mann, 2007, Ference and Barolo, 2006, Horabin, 2005, Rives et al., 2006, Wei et al., 2005, Chao et al., 2007, Gallet et al., 2006, Colomb et al., 2007, Ogden et al., 2006, de Navas et al., 2006, Karcavich and Doe, 2005, Strigini et al., 2006, Ayers et al., 2009, Song and Taylor, 2003, Meyer et al., 2006, Jennings et al., 2006, Takacs et al., 2008, Sasamura et al., 2007, Mayor et al., 2000, Han et al., 2000, Mellerick and Liu, 2004, Overton et al., 2007, Callejo et al., 2006, Smelkinson et al., 2007, López-Onieva et al., 2008, Estella and Mann, 2008, Lebreton et al., 2008, Wheeler et al., 2008, Tran and Doe, 2008, Thomsen et al., 2010, Chandraratna et al., 2007, Mitchell et al., 2008, Vincent et al., 2008, Peterson-Nedry et al., 2008, McKay et al., 2009, Dawber et al., 2005, Noyes et al., 2008, Smith-Bolton et al., 2009, Vied and Kalderon, 2009, Makhijani et al., 2007, Gambetta et al., 2009, Brodu et al., 2004, Seibert et al., 2009, Pines et al., 2010, Guo and Wang, 2009, Kim et al., 2007, Molnar et al., 2007, Bhatia et al., 2010, Wojcinski et al., 2011, Wang et al., 2011) EN (Ward et al., 2006, Takaesu et al., 2005, Zhao et al., 2007, Pallavi and Shashidhara, 2005, Mugat et al., 2008, Bury et al., 2008, Giang et al., 2011) en1 (The modENCODE Consortium, 2010, The modENCODE Consortium, 2010) Eng (Drewell et al., 2002, Nanda and Brand, 2000) Engrailed (Kang et al., 2006) Engrailed/Invected (Sedaghat et al., 2002) EnR (Dasen, 2005) Es   spa2 (Weyers et al., 2011) spt (Hadorn and Graber, 1946, ) V
Name Synonym	Apigmented abdomen   engrailed (Kategaya et al., 2009, Chen and Rasmuson-Lestander, 2009, Schwartz and Pirrotta, 2007, Hendriksen et al., 2005, Wang and Gergen, 2007, Crickmore and Mann, 2007, Casso et al., 2007, Mulinari et al., 2007, Jaeger and Reinitz, 2006, Tucker and Chiquet-Ehrismann, 2006, Wheeler et al., 2006, Beckett and Baylies, 2006, Bowler et al., 2006, Estes et al., 2008, Kremer et al., 2010, Park et al., 2008, Brown et al., 2005, Peel et al., 2006, Crickmore and Mann, 2007, Salazar-Ciudad, 2006, Muller and Kassis, 2006, Wendler et al., 2006, Casso et al., 2008, Anderson and Pick, 2008, Durant and Kassis, 2008, Garaulet and Sánchez-Herrero, 2008, Mulinari et al., 2008, Bejarano et al., 2005, Maeda et al., 2007, Staudt et al., 2006, Torroja et al., 2005, Shen et al., 2008, Jamieson et al., 2006, Wang et al., 2007, Ringrose and Paro, 2007, Pereanu and Hartenstein, 2006, Goldstein et al., 2005, Vanderzwan-Butler et al., 2007, Joly et al., 2007, Bolivar et al., 2006, Lee et al., 2007, Nekrasov et al., 2007, Eugster et al., 2007, Gallet et al., 2008, Coiffier et al., 2008, DeVido et al., 2008, Haecker et al., 2008, Lebreton et al., 2008, Umemori et al., 2007, Gebelein and Mann, 2007, Liu et al., 2008, Claret et al., 2007, Mulinari et al., 2008, Kim et al., 2008, Kondo et al., 2007, Dekanty et al., 2005, Chandraratna et al., 2007, Siera and Cline, 2008, Dominguez-Gimenez et al., 2007, Oktaba et al., 2008, Gorfinkiel and Arias, 2007, Garaulet et al., 2008, Bejarano and Milán, 2009, Ji et al., 2009, Fang et al., 2009, Schaaf et al., 2009, Larsen et al., 2009, Millard and Martin, 2008, Mieszczanek et al., 2008, Kwon et al., 2009, Bosveld et al., 2008, Mosimann et al., 2009, Gambetta et al., 2009, Zhai et al., 2009, Tolhuis et al., 2006, Tron et al., 2004, Hou et al., 2009, Mulinari and Häcker, 2009, Kumar et al., 2009, Cheng et al., 2010, Beckervordersandforth et al., 2008, Grieder et al., 2009, Landsberg et al., 2009, Eivers et al., 2009, Bischoff et al., 2009, Wilkie et al., 2001, Braid et al., 2010, Hartmann et al., 2010, Hsouna and VanBerkum, 2008, Blagburn, 2008, Gettings et al., 2010, Robinett et al., 2010, Xu et al., 2008, Wang et al., 2010, Jones et al., 2010, Ding et al., 2010, Lundell et al., 1996, Menéndez et al., 2010, Cunningham et al., 2010, Schwartz et al., 2010, Bhatia et al., 2010, Jung et al., 2010, Lee et al., 1999, Klebes et al., 2002, Fiedler et al., 2011, Hogan et al., 2011, Seibert and Urbach, 2010, Weyers et al., 2011, Sprecher and Hirth, 2006, Rand et al., 2010, McKnight et al., 2011) Engrailed (Goodman et al., 2006, Ward et al., 2006, Bajard, 2006, Noro and Mann, 2007, Ference and Barolo, 2006, Fraser, 2006, Herranz and Milan, 2006, Mahr and Aberle, 2006, Gallet et al., 2006, Bergsland et al., 2006, Tountas and Fortini, 2007, Cheesman et al., 2004, Karcavich and Doe, 2005, Strigini et al., 2006, Song and Taylor, 2003, Hsouna et al., 2003, McDonald et al., 2003, Meyer et al., 2006, Mayor et al., 2003, Molnar et al., 2011, Momota et al., 2011, Takacs et al., 2008, Sasamura et al., 2007, Religa et al., 2005, Von Ohlen et al., 2007, Israeli et al., 2007, Reig et al., 2007, Nanda and Brand, 2000, Han et al., 2000, Mellerick and Liu, 2004, Wawersik et al., 2005, Smelkinson et al., 2007, Ruel et al., 2007, Tran and Doe, 2008, Pallavi and Shashidhara, 2005, Berger et al., 2008, Bayraktar et al., 2006, Thomsen et al., 2010, Peterson-Nedry et al., 2008, Gaziova and Bhat, 2009, Orgogozo and Grueber, 2005, Dawber et al., 2005, Bossuyt et al., 2009, Franch-Marro et al., 2008, Mirth et al., 2009, Vied and Kalderon, 2009, Bejarano et al., 2007, Leal et al., 2009, Mugat et al., 2008, Liu et al., 2009, Pines et al., 2010, Bahri et al., 2001, Guo and Wang, 2009, Colomb et al., 2008, Rousset et al., 2010, Kopytova et al., 2006, Kim et al., 2007, Bury et al., 2008, Molnar et al., 2007, Tamori et al., 2010, Reddy et al., 2010, Emmons et al., 2007, Michaud and Tanguay, 2003, Djiane et al., 2011, Giang et al., 2011, Slack et al., 2006, Wang et al., 2011) Erased   Invected (Beckervordersandforth et al., 2008) sparse SGPs 2 (Weyers et al., 2011) spermatheca (Hadorn and Graber, 1946, ) transcript group V
Secondary FlyBase IDs
	FBgn0003492 FBgn0014157 FBgn0016969
References ( 1626 )
Generate a list of	All references relating to this gene ( 1626 )
List References by type	Research paper  ( 1116 ) Supplementary material  ( 12 ) Review  ( 308 ) Erratum  ( 2 ) Personal communication to FlyBase  ( 8 ) Abstract  ( 155 ) FlyBase analysis  ( 1 ) Conference report  ( 3 ) Stock list  ( 1 ) Book  ( 2 ) Protein sequence record  ( 1 ) Letter  ( 6 ) Computer file  ( 2 ) Biography  ( 1 ) Automatic genome annotation  ( 1 ) Curated genome annotation  ( 1 ) Book review  ( 1 ) Thesis  ( 1 ) Poem  ( 1 ) Interview  ( 2 )
Recent research papers ( 52 )
	Babcock et al., 2011, Curr. Biol. 21(18): 1525--1533 Hedgehog signaling regulates nociceptive sensitization. [FBrf0216292] Djiane et al., 2011, J. Cell Biol. 192(1): 189--200 Su(dx) E3 ubiquitin ligase-dependent and -independent functions of Polychaetoid, the Drosophila ZO-1 homologue. [FBrf0212737] Fiedler et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(5): 1937--1942 Dishevelled interacts with the DIX domain polymerization interface of Axin to interfere with its function in down-regulating {beta}-catenin. [FBrf0212953] Gabilondo et al., 2011, Mech. Dev. 128(3-4): 208--221 A targeted genetic screen identifies crucial players in the specification of the Drosophila abdominal Capaergic neurons. [FBrf0213290] Giang et al., 2011, J. Neurogenet. 25(1-2): 17--26 The Serotonin Transporter Expression in Drosophila melanogaster. [FBrf0213697] Hogan et al., 2011, PLoS Genet. 7(2): e1001305 Two frizzled planar cell polarity signals in the Drosophila wing are differentially organized by the fat/dachsous pathway. [FBrf0213159] McKnight et al., 2011, Mol. Cell. Biol. 31(23): 4746--4759 Extranucleosomal DNA binding directs nucleosome sliding by chd1. [FBrf0216578] Molnar et al., 2011, PLoS Genet. 7(3): e1001335 Role of the Drosophila non-visual ß-arrestin kurtz in hedgehog signalling. [FBrf0213301] Momota et al., 2011, Matrix Biol. 30(4): 258--266 Drosophila type XV/XVIII collagen, Mp, is involved in Wingless distribution. [FBrf0213898] Parker et al., 2011, Sci. Signal. 4(176): ra38 The cis-Regulatory Logic of Hedgehog Gradient Responses: Key Roles for Gli Binding Affinity, Competition, and Cooperativity. [FBrf0213871] Stagg et al., 2011, Development 138(11): 2171--2183 Dual role for Drosophila lethal of scute in CNS midline precursor formation and dopaminergic neuron and motoneuron cell fate. [FBrf0213671] Terriente-Félix et al., 2011, Dev. Biol. 350(2): 382--392 A conserved function of the chromatin ATPase Kismet in the regulation of hedgehog expression. [FBrf0212899] Walrad et al., 2011, Mol. Biol. Cell 22(8): 1364--1374 Hairless is a cofactor for Runt-dependent transcriptional regulation. [FBrf0213441] Wang et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(27): 11139--11144 Sexually dimorphic regulation of the Wingless morphogen controls sex-specific segment number in Drosophila. [FBrf0214239] Watson et al., 2011, Development 138(7): 1285--1295 Drosophila hedgehog signaling and engrailed-runt mutual repression direct midline glia to alternative ensheathing and non-ensheathing fates. [FBrf0213237] Weyers et al., 2011, Dev. Biol. 353(2): 217--228 A genetic screen for mutations affecting gonad formation in Drosophila reveals a role for the slit/robo pathway. [FBrf0213539] Wojcinski et al., 2011, Dev. Biol. 358(1): 168--180 DSulfatase-1 fine-tunes Hedgehog patterning activity through a novel regulatory feedback loop. [FBrf0215253] Yorimitsu et al., 2011, Dev. Biol. 356(2): 598--607 Defective proventriculus specifies the ocellar region in the Drosophila head. [FBrf0214458] Benitez et al., 2010, Int. J. Dev. Biol. 54(4): 643--653 Both jnk and apoptosis pathways regulate growth and terminalia rotation during Drosophila genital disc development. [FBrf0210146] Bhatia et al., 2010, Genes Cells 15(7): 725--735 Chromatin remodeling protein INO80 has a role in regulation of homeotic gene expression in Drosophila. [FBrf0211277] Braid et al., 2010, Dev. Biol. 343(1-2): 178--189 Nemo phosphorylates Even-skipped and promotes Eve-mediated repression of odd-skipped in even parasegments during Drosophila embryogenesis. [FBrf0211008] Cheng et al., 2010, Dev. Biol. 337(1): 99--109 Regulation of smoothened by Drosophila G-protein-coupled receptor kinases. [FBrf0209432] Chou et al., 2010, Cell 142(6): 954--966 Patterning axon targeting of olfactory receptor neurons by coupled hedgehog signaling at two distinct steps. [FBrf0211838] Cunningham et al., 2010, Mol. Cell. Biol. 30(3): 820--828 Characterization of the polycomb group response elements of the Drosophila melanogaster invected Locus. [FBrf0209731] Ding et al., 2010, Genetics 184(3): 717--729 Drosophila Raf's N terminus contains a novel conserved region and can contribute to torso RTK signaling. [FBrf0210411] Fabre et al., 2010, Development 137(17): 2885--2894 Mechanosensilla in the adult abdomen of Drosophila: engrailed and slit help to corral the peripheral sensory axons into segmental bundles. [FBrf0211534] Gettings et al., 2010, PLoS Biol. 8(6): e1000390 JNK Signalling Controls Remodelling of the Segment Boundary through Cell Reprogramming during Drosophila Morphogenesis. [FBrf0211014] Hartmann et al., 2010, Dev. Biol. 340(1): 125--133 Coral emx-Am can substitute for Drosophila empty spiracles function in head, but not brain development. [FBrf0210189] Jia et al., 2010, J. Biol. Chem. 285(48): 37218--37226 Casein kinase 2 promotes Hedgehog signaling by regulating both smoothened and Cubitus interruptus. [FBrf0214177] Jones et al., 2010, J. Cell Sci. 123(13): 2179--2189 Cytokinesis proteins Tum and Pav have a nuclear role in Wnt regulation. [FBrf0211056] Jung et al., 2010, Genomics 96(3): 154--166 Identification of conserved Drosophila-specific euchromatin-restricted non-coding sequence motifs. [FBrf0211521] Kannan et al., 2010, Mech. Dev. 127(1-2): 137--145 Abdominal-A mediated repression of Cyclin E expression during cell-fate specification in the Drosophila central nervous system. [FBrf0209859] Kazemian et al., 2010, PLoS Biol. 8(8): Quantitative analysis of the Drosophila segmentation regulatory network using pattern generating potentials. [FBrf0211688] Klein et al., 2010, Dev. Biol. 337(2): 458--470 The proprioceptive and contractile systems in Drosophila are both patterned by the EGR family transcription factor Stripe. [FBrf0209777] Kremer et al., 2010, Dev. Biol. 337(1): 110--123 In vivo analysis in Drosophila reveals differential requirements of contact residues in Axin for interactions with GSK3beta or beta-catenin. [FBrf0209417] Menéndez et al., 2010, Proc. Natl. Acad. Sci. U.S.A. 107(33): 14651--14656 A tumor-suppressing mechanism in Drosophila involving cell competition and the Hippo pathway. [FBrf0211593] Monier et al., 2010, Nat. Cell Biol. 12(1): 60--65 An actomyosin-based barrier inhibits cell mixing at compartmental boundaries in Drosophila embryos. [FBrf0209576] Pines et al., 2010, Development 137(6): 913--922 The cytolinker Pigs is a direct target and a negative regulator of Notch signalling. [FBrf0210124] Rand et al., 2010, Insect Biochem. Mol. Biol. 40(11): 792--804 Permeabilization of Drosophila embryos for introduction of small molecules. [FBrf0211955] Reddy et al., 2010, Mol. Cell. Biol. 30(21): 5234--5244 Drosophila Transcription Factor Tramtrack69 Binds MEP1 To Recruit the Chromatin Remodeler NuRD. [FBrf0212010] Robinett et al., 2010, PLoS Biol. 8(5): e1000365 Sex and the single cell. II. There is a time and place for sex. [FBrf0210735] Rousset et al., 2010, Development 137(13): 2177--2186 The Drosophila serine protease homologue Scarface regulates JNK signalling in a negative-feedback loop during epithelial morphogenesis. [FBrf0211045] Schubiger et al., 2010, Dev. Biol. 347(2): 315--324 Regeneration and transdetermination: The role of wingless and its regulation. [FBrf0212048] Schwartz et al., 2010, PLoS Genet. 6(1): e1000805 Alternative epigenetic chromatin states of polycomb target genes. [FBrf0209708] Seibert and Urbach, 2010, Dev. Biol. 346(2): 332--345 Role of en and novel interactions between msh, ind, and vnd in dorsoventral patterning of the Drosophila brain and ventral nerve cord. [FBrf0211900] Subramanian and Gadgil, 2010, IET Syst. Biol. 4(2): 169 Robustness of the Drosophila segment polarity network to transient perturbations. [FBrf0210287] Tamori et al., 2010, PLoS Biol. 8(7): e1000422 Involvement of Lgl and Mahjong/VprBP in Cell Competition. [FBrf0211298] Thomsen et al., 2010, Development 137(17): 2951--2960 Developmental RNA processing of 3'UTRs in Hox mRNAs as a context-dependent mechanism modulating visibility to microRNAs. [FBrf0211493] Walrad et al., 2010, Mol. Biol. Cell 21(13): 2315--2326 Distinct contributions of conserved modules to runt transcription factor activity. [FBrf0211193] Wang et al., 2010, Dev. Biol. 337(2): 246--258 The role of eyg Pax gene in the development of the head vertex in Drosophila. [FBrf0209801] Wang et al., 2010, PLoS ONE 5(7): e11498 NELF Potentiates Gene Transcription in the Drosophila Embryo. [FBrf0211281] Yassin et al., 2010, Evol. Dev. 12(3): 288--295 Catching the phylogenic history through the ontogenic hourglass: a phylogenomic analysis of Drosophila body segmentation genes. [FBrf0211054] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...