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Gene Dmel\dl

General Information
Symbol	Dmel\dl	Species	D. melanogaster
Name	dorsal	Annotation symbol	CG6667
Feature type	protein_coding_gene	FlyBase ID	FBgn0260632
Gene Model Status	Current	Stock availability	20 publicly available
Genomic Location
Chromosome (arm)	2L	Recombination map	2-52.9
Cytogenetic map	36C8-36C9	Sequence location	2L:17,436,830..17,450,360 [-]
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 dorsal is referred to in FlyBase by the symbol Dmel\dl (CG6667, FBgn0260632). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: RNA polymerase II regulatory region sequence-specific DNA binding; protein binding; sequence-specific DNA binding; RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in positive regulation of transcription; sequence-specific distal enhancer binding RNA polymerase II transcription factor activity; high mobility group box 1 binding; repressing transcription factor binding. There is experimental evidence that it is involved in the biological process: immune response; melanization defense response; peripheral nervous system neuron development; positive regulation of transcription from RNA polymerase II promoter; negative regulation of gene expression; maternal determination of anterior/posterior axis, embryo; regulation of hemocyte proliferation; regulation of alternative nuclear mRNA splicing, via spliceosome; plasmatocyte differentiation. 156 alleles are reported. The phenotypes of these alleles are annotated with: organ system subdivision; organ system; adult segment; portion of tissue; multicellular structure; extended germ band embryo; imaginal precursor; epithelial furrow; spiracle; external compound sense organ. It has 5 annotated transcripts and 5 annotated polypeptides. Protein features are: Cell surface receptor IPT/TIG; Immunoglobulin E-set; Immunoglobulin-like fold; NF-kappa-B/Rel/dorsal; Rel homology; p53-like transcription factor, DNA-binding. 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 00-06 and 18-24 hour embryonic stages. Summary of FlyAtlas Anatomical Expression Data: Two or more Affy2 ProbeSets identify exons of this gene. This is a summary of the tissue expression peaks exhibited in at least one of these ProbeSets. Expression at high levels in the following post-embryonic organs or tissues: adult crop, adult salivary gland. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, larval/adult midgut, larval/adult hindgut, adult heart, larval salivary gland, larval trachea, adult ovary, larval/adult carcass. Gene sequence location is 2L:17436830..17450360.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	dl: dorsal Embryos produced by homozygous dl females form normal cellular blastoderm but at gastrulation develop into yolk-filled tube of dorsal hypoderm. Hair pattern of cuticle characteristic of dorsal hypoderm; ventral structures, such as denticle belts, lacking. Normally, dorsal infoldings occupy entire circumference of embryo. Evidence of anterior-posterior differentiation includes possible mouth armature structures anteriorly, small spiracles posteriorly, and orientation of hairs. The periodicity of stripes of ftz expression in pre gastrulation embryos, as revealed by antibody staining, displays the pattern normally characteristic of the dorsum circumferentially in embryos produced by dl females (Carroll, Winslow, Twombly, and Scott, 1987, Development 99: 327-32. Embryos produced by dl/dl and dl/Df(2L)TW137 indistinguishable, suggesting dl to be amorphic. Penetrance complete; expression constant. Embryos of dl2 females lack all structures normally derived from the ventral half of the egg, including mesoderm, endodermal gut, ventral nervous system, and ventral hypoderm. dl1 and to a lesser extent dl2, females produce embryos with reduced capacity for neurogenesis in response to an absence of dl function (Campos-Ortega, 1983, Wilhelm Roux's Arch. Dev. Biol. 192: 317-26). dl germ line dependent; homozygous germ-line clones produce dorsalized embryos (Schupbach and Wieschaus, 1986, Dev. Biol. 113: 443-48). Embryos of dl/+ females produced at 29 develop into comparatively normal-looking larvae; they mainly lack internal organs, such as mesoderm and parts of the anterior and posterior gut; often ventral hypoderm including denticle belts reduced; phenotype sensitive to genetic background. At 22, dl/+ females produce normal embryos. Developmental fate of ventrally located cells on cellular blastoderm apparently shifted to that of more dorsally located cells. The phenotype of embryos produced by dl/dl females partially rescued by the injection of wild-type cytoplasm but not RNA (Santamaria and Nusslein-Volhard, 1983, EMBO J. 2: 1695-99; Anderson and Nusslein-Volhard, 1944, Nature 34: 225-27). Developmental profiles show transcript to be present only in ovaries and pre-cellular-blastoderm stages of embryogenesis. In situ hybridization indicates that ovarian transcript accumulates in nurse cells from stage 5 to 11; number of transcripts per genome equivalent in these polytene cells remains low and constant until stage 10, at which time there is a dramatic increase in the relative numbers of transcripts. After a lag of one or two nuclear divisions, transcript begins to accumulate in the oocyte; by stage 12 there is little detectable transcript in the nurse cells. It appears as though the nurse-cell transcript is transferred to the oocyte and thus to the embryo; transcript seems to be uniformly distributed in stage 14 oocytes (Steward, Ambrosio, and Schedl). dl protein is uniformly distributed throughout cytoplasm of early embryo; in the syncytial blastoderm a gradient of expression is achieved by the graded transport of dl protein into nuclei, with the highest nuclear concentrations found ventrally; protein remains cytoplasmic dorsally. Maternal dorsalizing mutants prevent nuclear localization and ventralized embryos show dorsal as well as ventral nuclear localization (Steward, Zusman, Huang, and Schedl, 1988, Cell 55: 487-95; Rushlow, Han, Manley, and Levine, 1989, Cell 59: 1165-77; Steward, 1989, Cell 59: 11179-88; Roth, Stein, and Nusslein-Volhard, 1989, Cell 59: 1189-1202).
Recent Updates
Description	What does this section display? This section contains items that were added to this record for each release. It currently only tracks new links between this FlyBase report and other FlyBase data classes (e.g. genes, references, stocks) or controlled vocabulary terms (e.g. GO, anatomy terms). What does this section not display? This section does not currently display links that were removed or gene model changes.
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FB2011_10	References Kechris and Li, 2008, BMC Bioinformatics 9: 506
FB2012_01	Insertions Mi{MIC}dl[MI03808] Sequence features HFA03521 BKN27820 HFA03520 References Benveniste et al., 1998, Development 125(23): 4757--4765 Rand et al., 2010, Insect Biochem. Mol. Biol. 40(11): 792--804 Negreiros et al., 2010, genesis 48(1): 31--43 Bellen, 2011.10.25, set956.xml Alleles dl[MI03808] Controlled Vocabulary Terms wandering third instar larval stage neuromuscular junction third instar larval stage L1 negative regulation of gene expression RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in positive regulation of transcription sequence-specific distal enhancer binding RNA polymerase II transcription factor activity prepupal stage P3 subsynaptic reticulum prepupal stage P1 type I bouton third instar larval stage L2 embryonic/larval hypodermal muscle A1-7 ventral longitudinal muscle 4 postsynaptic membrane A1-7 ventral longitudinal muscle 3 prepupal stage P2 RNA polymerase II regulatory region sequence-specific DNA binding
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 36C8-36C9   Limits computationally determined from genome sequence between P{lacW}Mhck10423&P{lacW}Cask03902 and P{lacW}Aac11k06710
Experimentally Determined Cytological Location
Cytogenetic map Notes References 36C9-36C9   Location inferred from insertion in: dl[GSd447] ; Limits computationally determined from genome sequence between P{lacW}Mhck10423&P{lacW}Cask03902 and P{lacW}Aac11k06710   (Minidorff et al., 2007) 36C-36C   (determined by in situ hybridisation)   (Spradling et al., 1999) 36D1-36D3     (BDGP Project Members, 1994-1999) 36C-36C   (determined by in situ hybridisation)   (Steward et al., 1984)
Experimentally Determined Recombination Data
Location	2-52.9 (Tearle and Nusslein-Volhard, 1987) 2-53 (Schupbach and Wieschaus, 1989)
Left of (cM)	nub (Szabad et al., 1989)
Right of (cM)	al (Szabad et al., 1989)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\dl 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) dl-RA FBtr0081005  NM_165217   2485   677 dl-RB FBtr0081006  NM_165218   2828   677 dl-RC FBtr0081007  NM_165219   4767   999 dl-RD FBtr0301383  NM_001169529   2590   677 dl-RE FBtr0301384  NM_001169530   2504   677
Additional Transcript Data & Comments
Reported size (kB)	4.4, 2.8 (northern blot) (Gross et al., 1999) 2.8 (northern blot) (Steward et al., 1984) 2.8 (longest cDNA) (Steward, 1987)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein dl-PA   FBpp0080558   75.3   677   8.10     NP_724052   AAF53611 dl-PB   FBpp0080559   75.3   677   8.10     NP_724053   AAF53610 dl-PC   FBpp0080560   111.6   999   5.04     NP_724054   AAF53612 dl-PD   FBpp0290597   75.3   677   8.10     NP_001163000   ACZ94286 dl-PE   FBpp0290598   75.3   677   8.10     NP_001163001   ACZ94287
Additional Polypeptide Data & Comments
Reported size (kDa)	994 (aa); 190 (kD observed); 110 (kD predicted) (Gross et al., 1999) 677 (aa); 75.6 (kD predicted) (Steward, 1987) 85 (kD observed) (Roth et al., 1989) 85 (kD) (Govind et al., 1992) 83 (kD observed) (Steward, 1989) 75 (kD predicted) (Steward et al., 1988)
Comments	lwr protein was shown to bind and conjugate smt3 protein to dl protein. In addition, lwr protein releives the inhibition of dl protein nuclear uptake by cact protein in cultured cells. (Bhaskar et al., 2000) The unique C-terminal portion of the large dl protein isoform has a transactivation domain and is able to induce the expression of a reporter gene in transient transfection assays in mouse 3T3 cells. The C-terminal activation domain was mapped by deletion mapping to between residues 576 and 638. (Gross et al., 1999) Two mutant dl proteins (dlS234P.Scer\adh1 and dlC233R.Scer\adh1) that activate transcription but are insensitive to inhibition by cact were isolated by a yeast assay. These mutant dl proteins are able to enter the nucleus, bind DNA, and activate transcription but are unable to bind cact protein. The surface of dl protein that is likely to bind cact protein was identified. (Lehming et al., 1995) Two mutant dl proteins (dlS234P.Scer\adh1 and dlC233R.Scer\adh1) that activate transcription but are insensitive to inhibition by cact were isolated by a yeast assay. These mutant dl proteins are able to enter the nucleus, bind DNA, and activate transcription but are unable to bind cact protein. The surface of dl protein that is likely to bind cact protein was identified. (Lehming et al., 1995) Small sequential deletions in the dl protein were used to map various functional regions of the protein. Sequences that are necessary for DNA binding map between residues 47 and 230. Two regions were mapped which are involved with inhibitor binding. Loss of sequences between residues 218-245 (region I) or residues 322-241 (region II)abolishes binding to cact protein. Deletions in residues 246-321 (between regions I and II) reduce but do not abolish cact binding. Mutation of highly conserved residues 231-237 in region I nearly abolishes binding. Mutation of sequences in region II containing the nuclear localization signal (residues 335-340) result in near or complete failure to bind cact protein. Finally, point mutations in the region I sequence can uncouple DNA binding and inhibitor interactions. (Tatei and Levine, 1995) Disulfide crosslinking studies were used to show that dl and cact proteins exist as three different complexes in the embyro. Complex 1 (190kD) is a dl homodimer (dl2). Complex 2 (270kD)consists of a complex 1 and a cact molecule (dl2cact). Complex 3 is a cact protein complex. In wild type embryos, complex 1 was observed as the major form of dl protein and complex 2 was a minor form. Virtually no dl monomer was detected. Mutant analysis indicates that complex 1 is a cytoplasmic form while complex 2 is mainly nuclear. (Isoda and Nusslein-Volhard, 1994) Immunoprecipitation studies in embryos which contain a dl-lacZ protein fusion construct show that the dl protein can self associate in a protein complex. In early embryonic extracts, dl protein is found in large complexes of about 200kD. These are thought to be either dl homodimers and cact protein or dl protein multimers. (Govind et al., 1992) The cact binding site on the dl protein was localized to the region between amino acids 168 and 350 and is thought to include residues around aa270. This is within the RH domain (amino acid 47-341) and overlaps with or is adjacent to the nuclear localization signal. It overlaps with a region shown to be necessary for dl protein to be retained in the cytoplasm (FBrf0056112). (Kidd, 1992) Cotransfection experiments in Schneider cells show that dl protein activates CAT expression from a zen-CAT construct. CAT activity is enhanced in the presence of either Tl or Pka-C1 proteins. Both Tl and Pka-C1 proteins affect dl protein nuclear localization and dl protein activity in the nucleus. Evidence suggests that the signalling pathway from Tl to dl proteins acts through Pka-C1 protein. dl mutant constructs were tested to map the regions of dl protein required for responses to Tl and Pka-C1 proteins. (Norris and Manley, 1992)
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • NF-kappa-B/Rel/dorsal (IPR000451) Cell surface receptor IPT/TIG (IPR002909) p53-like transcription factor, DNA-binding (IPR008967) Rel homology (IPR011539) Immunoglobulin-like fold (IPR013783) Immunoglobulin E-set (IPR014756)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA201313   AA538942   AE014134 AAF53610   AE014134 AAF53611   AE014134 AAF53612   AE014134 ACZ94286   AE014134 ACZ94287   AF053614 AAC35296   AI108304   AI546279   AQ025943   BF502724   BH900897   BI216660   BI367180   BI370432   BI372084   BI372419   BI482551   BI571425   BI640385   BP540348   BP540349   BP545366   BP545367   BP559892   BP559893   BT015205 AAT94434   BT099643   CO155544   CO298488   CO309086   CO309911   CO312363   CZ469985   CZ474527   CZ476269   EC077776   EC086724   EC087684   EC202035   EC233251   EC235564   EC247891   EV585118   GH943943   GH946577   GH946884   GH953375   GH958575   GH959097   GH960664   GH963271   GH964582   GH965825   GH966137   M23702 AAA28479   X65272
UniProtKB/Swiss-Prot	P15330
UniProtKB/TrEMBL
Mapped Features
Mapped Features have been reorganized, please see this article for details. Additional mapped features and mutations can be found on GBrowse or related reports. Type Symbol & Location Additional Notes References
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage 1 -- 3   organism | ubiquitous   (Steward et al., 1988) northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage 1 -- 5     (Steward et al., 1984) Comment:reference states 0-2.5 hr AEL embryonic stage 11 -- 17     (Gross et al., 1999) Comment:reference states >=6 hr AEL larval stage     (Gross et al., 1999) embryonic/larval fat body   (Gross et al., 1999) adult stage | female     (Gross et al., 1999) adult stage | male     (Gross et al., 1999) adult stage & oogenesis   ovary   (Steward et al., 1984)
Additional Descriptive Data
	The larger form of dl transcript is expressed in a tissue specific manner. It is present in larvae and adults of both sexes. The intensity of the band increases upon immune challenge especially in larvae and adult males. dl transcripts are present at low levels in the gut and fat body of unchallenged larvae. The levels are enhanced in the larval fat body after immune challenge. In embryos the larger dl transcript is present from 6-9hr embryos on through embryogenesis. (Gross et al., 1999) dl transcripts are detected uniformly in embryos up until the cellular blastoderm stage. (Steward et al., 1988)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
No Assay Recorded Stage Tissue/Position (including subcellular localization) Reference embryonic stage 2 -- 4   organism | dorsal ventral gradient   (Rushlow et al., 1989) embryonic stage 3 -- 5   organism | dorsal ventral gradient   (Roth et al., 1989) embryonic stage 3 -- 7   organism | dorsal ventral gradient   (Steward et al., 1988) embryonic stage 7 -- 14   ventral furrow   (Rushlow et al., 1989) immunolocalization Stage Tissue/Position (including subcellular localization) Reference embryonic stage 2   organism | ubiquitous   (Roth et al., 1989) embryonic stage 2 -- 3   organism | ubiquitous   (Steward et al., 1988) embryonic stage 3 -- 9   organism | dorsal ventral gradient   (Steward, 1989) embryonic stage 4 -- 5   organism | dorsal   (Sun et al., 2004) embryonic stage 4 -- 7   organism | dorsal   (Rao et al., 1991) embryonic stage 7   mesoderm   (Roth et al., 1989) embryonic stage 7 -- 14   mesoderm   (Rushlow et al., 1989) wandering third instar larval stage   embryonic/larval hypodermal muscle   (Cantera et al., 1999) A1-7 ventral longitudinal muscle 3   • neuromuscular junction (Bolatto et al., 2003) A1-7 ventral longitudinal muscle 4   • neuromuscular junction (Bolatto et al., 2003) wandering third instar larval stage -- prepupal stage P3   type I bouton   • subsynaptic reticulum (Cantera et al., 1999) adult stage   adult neuromuscular junction   (Beramendi et al., 2007) adult stageoogenesis stage S9 -- S12   nurse cell   (Rushlow et al., 1989) mass spectroscopy Stage Tissue/Position (including subcellular localization) Reference day 7 of adulthood -- day 49 of adulthood   adult heart   (Cammarato et al., 2011) western blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage 1 -- 17     (Steward, 1989) adult stage   ovary   (Steward, 1989) unfertilized egg stage   unfertilized egg   (Steward, 1989)
Additional Descriptive Data
	Up until embryonic stage 3, dl protein is localized primarily in the cytoplasm. Between stages 3 and 4 dl protein is localized to nuclei along the ventral side of the embryo and persists through grastrulation. (Roth et al., 1989, Steward et al., 1988) dl protein is cytoplasmic until embryonic cleavage cycle 10, at which point it becomes nuclear. A specific nuclear transport system is proposed to be required for wild type dl function. (Steward et al., 1988, Rushlow et al., 1989) The larger form of dl protein translocates into the nucleus after bacterial challenge. (Beramendi et al., 2007, Gross et al., 1999) cact colocalizes with Fas2 at the larval neuromuscular junction, but is shifted more cytoplasmically than is Fas2. (Bolatto et al., 2003) The dl protein is expressed throughout the embryo, however, by stage 2 markedly higher levels of staining are detected in the ventral region, which persists through gastrulation. dl protein is located primarily in the cytoplasm of nurse cells, and the embryo and moves into the ventral nuclei at cleavage cycle 11. This transition to nuclear localization is proposed to be required for dl function, as it is not observed in dl mutants. (Rushlow et al., 1989) dl protein is diffusely distributed in the nucleus and cytoplasm of somatic muscles during the last hours of larval development, and though the first four hours of pupariation. dl protein is enriched in the subsynaptic reticulum of type I synaptic boutons. Four hours after pupariation, dl protein is shifted to the nucleus. In adults, the dl protein distibution is similar to that in larvae. (Cantera et al., 1999)
Marker for
Subcellular Localization
CV Term	nucleus (Steward, 1989) nucleus cytoplasm (Rushlow et al., 1989) cytoplasm nucleus (Roth et al., 1989) nucleus (Gross et al., 1999)
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 FBgn0260632    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0260632 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 00-06 and 18-24 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 FBgn0260632 expression level Developmental Stage   Expression Level embryo 00-02hr    1827 embryo 02-04hr    107 embryo 04-06hr    32 embryo 06-08hr    28 embryo 08-10hr    60 embryo 10-12hr    171 embryo 12-14hr    420 embryo 14-16hr    929 embryo 16-18hr    1185 embryo 18-20hr    1441 embryo 20-22hr    1616 embryo 22-24hr    1746 larva L1    829 larva L2    713 larva L3 12hr old    615 larva L3 puffstage 1-2    579 larva L3 puffstage 3-6    719 larva L3 puffstage 7-9    619 white prepupae new    611 white prepupae 12hr    861 white prepupae 24hr    533 pupae 2d postWPP    1057 pupae 3d postWPP    1054 pupae 4d postWPP    1024 adult male 01day    826 adult male 05day    950 adult male 30day    1104 adult female 01day    942 adult female 05day    1138 adult female 30day    1295 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  (1827) embryo 02-04hr  107 embryo 04-06hr    32 embryo 06-08hr    28 embryo 08-10hr    60 embryo 10-12hr  (171) embryo 12-14hr  (420) embryo 14-16hr  (929) embryo 16-18hr  (1185) embryo 18-20hr  (1441) embryo 20-22hr  (1616) embryo 22-24hr  (1746) larva L1  (829) larva L2  (713) larva L3 12hr old  (615) larva L3 puffstage 1-2  (579) larva L3 puffstage 3-6  (719) larva L3 puffstage 7-9  (619) white prepupae new  (611) white prepupae 12hr  (861) white prepupae 24hr  (533) pupae 2d postWPP  (1057) pupae 3d postWPP  (1054) pupae 4d postWPP  (1024) adult male 01day  (826) adult male 05day  (950) adult male 30day  (1104) adult female 01day  (942) adult female 05day  (1138) adult female 30day  (1295) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (1827) embryo 02-04hr    107 embryo 04-06hr    32 embryo 06-08hr    28 embryo 08-10hr    60 embryo 10-12hr    171 embryo 12-14hr    420 embryo 14-16hr    929 embryo 16-18hr    1185 embryo 18-20hr  1441 embryo 20-22hr  (1616) embryo 22-24hr  (1746) larva L1    829 larva L2    713 larva L3 12hr old    615 larva L3 puffstage 1-2    579 larva L3 puffstage 3-6    719 larva L3 puffstage 7-9    619 white prepupae new    611 white prepupae 12hr    861 white prepupae 24hr    533 pupae 2d postWPP    1057 pupae 3d postWPP    1054 pupae 4d postWPP    1024 adult male 01day    826 adult male 05day    950 adult male 30day    1104 adult female 01day    942 adult female 05day    1138 adult female 30day    1295 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    1827 embryo 02-04hr    107 embryo 04-06hr    32 embryo 06-08hr    28 embryo 08-10hr    60 embryo 10-12hr    171 embryo 12-14hr    420 embryo 14-16hr    929 embryo 16-18hr    1185 embryo 18-20hr    1441 embryo 20-22hr    1616 embryo 22-24hr    1746 larva L1    829 larva L2    713 larva L3 12hr old    615 larva L3 puffstage 1-2    579 larva L3 puffstage 3-6    719 larva L3 puffstage 7-9    619 white prepupae new    611 white prepupae 12hr    861 white prepupae 24hr    533 pupae 2d postWPP    1057 pupae 3d postWPP    1054 pupae 4d postWPP    1024 adult male 01day    826 adult male 05day    950 adult male 30day    1104 adult female 01day    942 adult female 05day    1138 adult female 30day    1295 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    1827 embryo 02-04hr    107 embryo 04-06hr    32 embryo 06-08hr    28 embryo 08-10hr    60 embryo 10-12hr    171 embryo 12-14hr    420 embryo 14-16hr    929 embryo 16-18hr    1185 embryo 18-20hr    1441 embryo 20-22hr    1616 embryo 22-24hr    1746 larva L1    829 larva L2    713 larva L3 12hr old    615 larva L3 puffstage 1-2    579 larva L3 puffstage 3-6    719 larva L3 puffstage 7-9    619 white prepupae new    611 white prepupae 12hr    861 white prepupae 24hr    533 pupae 2d postWPP    1057 pupae 3d postWPP    1054 pupae 4d postWPP    1024 adult male 01day    826 adult male 05day    950 adult male 30day    1104 adult female 01day    942 adult female 05day    1138 adult female 30day    1295 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0260632 expression level Developmental Stage   Expression Level embryo 00-02hr    1827 embryo 02-04hr    107 embryo 04-06hr    32 embryo 06-08hr    28 embryo 08-10hr    60 embryo 10-12hr    171 embryo 12-14hr    420 embryo 14-16hr    929 embryo 16-18hr    1185 embryo 18-20hr    1441 embryo 20-22hr    1616 embryo 22-24hr    1746 larva L1    829 larva L2    713 larva L3 12hr old    615 larva L3 puffstage 1-2    579 larva L3 puffstage 3-6    719 larva L3 puffstage 7-9    619 white prepupae new    611 white prepupae 12hr    861 white prepupae 24hr    533 pupae 2d postWPP    1057 pupae 3d postWPP    1054 pupae 4d postWPP    1024 adult male 01day    826 adult male 05day    950 adult male 30day    1104 adult female 01day    942 adult female 05day    1138 adult female 30day    1295 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (1827) embryo 02-04hr  107 embryo 04-06hr    32 embryo 06-08hr    28 embryo 08-10hr    60 embryo 10-12hr  (171) embryo 12-14hr  (420) embryo 14-16hr  (929) embryo 16-18hr  (1185) embryo 18-20hr  (1441) embryo 20-22hr  (1616) embryo 22-24hr  (1746) larva L1  (829) larva L2  (713) larva L3 12hr old  (615) larva L3 puffstage 1-2  (579) larva L3 puffstage 3-6  (719) larva L3 puffstage 7-9  (619) white prepupae new  (611) white prepupae 12hr  (861) white prepupae 24hr  (533) pupae 2d postWPP  (1057) pupae 3d postWPP  (1054) pupae 4d postWPP  (1024) adult male 01day  (826) adult male 05day  (950) adult male 30day  (1104) adult female 01day  (942) adult female 05day  (1138) adult female 30day  (1295) Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  1827 embryo 02-04hr    107 embryo 04-06hr    32 embryo 06-08hr    28 embryo 08-10hr    60 embryo 10-12hr    171 embryo 12-14hr    420 embryo 14-16hr    929 embryo 16-18hr    1185 embryo 18-20hr  1441 embryo 20-22hr  1616 embryo 22-24hr  1746 larva L1    829 larva L2    713 larva L3 12hr old    615 larva L3 puffstage 1-2    579 larva L3 puffstage 3-6    719 larva L3 puffstage 7-9    619 white prepupae new    611 white prepupae 12hr    861 white prepupae 24hr    533 pupae 2d postWPP    1057 pupae 3d postWPP    1054 pupae 4d postWPP    1024 adult male 01day    826 adult male 05day    950 adult male 30day    1104 adult female 01day    942 adult female 05day    1138 adult female 30day    1295 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    1827 embryo 02-04hr    107 embryo 04-06hr    32 embryo 06-08hr    28 embryo 08-10hr    60 embryo 10-12hr    171 embryo 12-14hr    420 embryo 14-16hr    929 embryo 16-18hr    1185 embryo 18-20hr    1441 embryo 20-22hr    1616 embryo 22-24hr    1746 larva L1    829 larva L2    713 larva L3 12hr old    615 larva L3 puffstage 1-2    579 larva L3 puffstage 3-6    719 larva L3 puffstage 7-9    619 white prepupae new    611 white prepupae 12hr    861 white prepupae 24hr    533 pupae 2d postWPP    1057 pupae 3d postWPP    1054 pupae 4d postWPP    1024 adult male 01day    826 adult male 05day    950 adult male 30day    1104 adult female 01day    942 adult female 05day    1138 adult female 30day    1295 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    1827 embryo 02-04hr    107 embryo 04-06hr    32 embryo 06-08hr    28 embryo 08-10hr    60 embryo 10-12hr    171 embryo 12-14hr    420 embryo 14-16hr    929 embryo 16-18hr    1185 embryo 18-20hr    1441 embryo 20-22hr    1616 embryo 22-24hr    1746 larva L1    829 larva L2    713 larva L3 12hr old    615 larva L3 puffstage 1-2    579 larva L3 puffstage 3-6    719 larva L3 puffstage 7-9    619 white prepupae new    611 white prepupae 12hr    861 white prepupae 24hr    533 pupae 2d postWPP    1057 pupae 3d postWPP    1054 pupae 4d postWPP    1024 adult male 01day    826 adult male 05day    950 adult male 30day    1104 adult female 01day    942 adult female 05day    1138 adult female 30day    1295 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 FBgn0260632    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0260632 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Two or more Affy2 ProbeSets identify exons of this gene. This is a summary of the tissue expression peaks exhibited in at least one of these ProbeSets. Expression at high levels in the following post-embryonic organs or tissues: adult crop, adult salivary gland. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, larval/adult midgut, larval/adult hindgut, adult heart, larval salivary gland, larval trachea, adult ovary, larval/adult carcass. [download data (TSV)] Guide to FlyAtlas expression level colors   No expression (0 - 9.999)   Low expression (10 - 99.999)   Moderate expression (100 - 499.999)   High level expression (500 - 999.999)   Very high expression (1000 - 25000) Heatmap Tissue   Expression Level Larval Central Nervous System no informative data Larval Midgut no informative data Larval Hindgut no informative data Larval Malpighian Tubules no informative data Larval Fat Body no informative data Larval Salivary Gland no informative data Larval Trachea no informative data Larval Carcass no informative data Adult Head no informative data Adult Eye no informative data Adult Brain no informative data Adult Thoracic-Abdominal Ganglion no informative data Adult Crop no informative data Adult Midgut no informative data Adult Hindgut no informative data Adult Malpighian Tubules no informative data Adult Fat Body     Adult Salivary Gland no informative data Adult Heart no informative data Adult VirginFemale Spermatheca no informative data Adult InseminatedFemale Spermatheca no informative data Adult Ovary no informative data Adult Testis no informative data Adult Male Accessory Gland no informative data Adult Carcass no informative data 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_10
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0260632
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal dl3 dl16 dl19 dl21 dl22 dl23 lethal, with Scer\GAL4hs.PB dlScer\UAS.cMa lethal, with Scer\GAL4scrt-11-6 dlGSd447 lethal | dominant | maternal effect dl24 dl25 dl26 dlmC1.Hsp83 lethal | embryonic stage | germline clone | non-rescuable maternal effect | recessive dl2L-136-13 dl2L-164-16 dl2L-258-38 dl2L-291-17 dl2L-294-9 dl2L-317-23 dl2L-325-31 dl2L-327-18 lethal | embryonic stage | maternal effect | recessive dl19 dl20 dl21 dl22 lethal | embryonic stage | maternal effect | recessive | heat sensitive dl17 dl18 lethal | embryonic stage | recessive dl1 lethal | first instar larval stage, with Scer\GAL4arm.PS dlScer\UAS.T:Avic\GFP-EGFP lethal | maternal effect dl5 dl6 dl7 dl8 lethal | maternal effect | heat sensitive dl2 dl17 lethal | maternal effect | recessive dl1 dl2 lethal | recessive dl38 lethal | second instar larval stage, with Scer\GAL4arm.PS dlScer\UAS.T:Avic\GFP-EGFP semi-lethal dlΔNLS viable, with Scer\GAL4Cg.PA dlScer\UAS.T:Avic\GFP-EGFP viable, with Scer\GAL4pnr-MD237 dlGD1238 Sterility Allele female sterile | dominant dl26 dl27 dlHsp83.T:Ecol\lacZ dlN.Hsp83.T:Ecol\lacZ female sterile | dominant | partially dlRHR.Hsp83.T:Ecol\lacZ female sterile | recessive dl17 dl18 dl19 dl20 dl21 dl22 dl01313 dlk10816 female sterile germline-dependent | dominant dl24 dl25 female sterile soma-dependent | dominant dl24 dl25 Other Phenotypes Allele immune response defective | conditional dl15 neuroanatomy defective, with Scer\GAL47B dlUY2278 dlUY3046 neuroanatomy defective, with Scer\GAL4ey-OK107 dlUY2278 neuroanatomy defective, with Scer\GAL4GMR.PF dlB.Scer\UAS.T:Ivir\HA1 dlGSd447 dlScer\UAS.cMa neuroanatomy defective, with Scer\GAL4MT14 dlGSd447 neuroanatomy defective | embryonic stage, with Scer\GAL4scrt-11-6 dlGSd447 visible, with Scer\GAL4bbg-C96 dld05894 visible, with Scer\GAL4GMR.PF dlB.Scer\UAS.T:Ivir\HA1 visible | cold sensitive (with dl1) dl4 visible | cold sensitive (with dl4) dl1 visible | dominant dl4 visible | temperature conditional, with Scer\GAL4sca-537.4 dlUY2278 wild-type dl+mWT dlbcd.PB dlHsp83.PG dlS79A dlT85A dlT290V dlY324F Phenotype manifest in Allele adult salivary gland primordium dl1 amnioproctodeal invagination dl1 axon & eye photoreceptor cell, with Scer\GAL4GMR.PF dlScer\UAS.cMa dlGSd447 dlB.Scer\UAS.T:Ivir\HA1 axon & eye photoreceptor cell, with Scer\GAL4MT14 dlGSd447 cephalopharyngeal skeleton | embryonic stage dl24 dl25 dl26 dl27 cephalopharyngeal skeleton | maternal effect dl1 dl17 dl18 dl19 dl20 dl21 dl22 denticle dl1 embryo dlΔNLS dl5 dlS234P dlunspecified embryo | ventral dlunspecified embryonic/first instar larval cuticle dl1 dl15 embryonic/first instar larval cuticle | maternal effect dl1 embryonic/first instar larval cuticle | maternal effect (with Df(2L)H20) dlC447 dlD447 embryonic/first instar larval cuticle | maternal effect | ventral dl17 dl18 dl19 dl20 dl21 dl22 embryonic/first instar larval cuticle | ventral dl1 dl15 dl2 embryonic epidermis | maternal effect dl1 embryonic epidermis | ventral dl16 dl19 dl21 dl22 dl23 dl3 endoderm dl1 eye, with Scer\GAL4GMR.PF dlB.Scer\UAS.T:Ivir\HA1 filzkorper dl1 filzkorper | maternal effect dl1 Kenyon cell, with Scer\GAL47B dlUY2278 dlUY3046 Kenyon cell, with Scer\GAL4ey-OK107 dlUY2278 lamina, with Scer\GAL4GMR.PF dlScer\UAS.cMa dlGSd447 dlB.Scer\UAS.T:Ivir\HA1 lamina, with Scer\GAL4lz-gal4 dlGSd447 macrochaeta dl4 macrochaeta | ectopic | cold sensitive (with dl1) dl4 macrochaeta | ectopic | cold sensitive (with dl4) dl1 macrochaeta | ectopic | temperature conditional, with Scer\GAL4sca-537.4 dlUY2278 macrochaeta | temperature conditional, with Scer\GAL4sca-537.4 dlUY2278 medulla, with Scer\GAL4GMR.PF dlScer\UAS.cMa dlGSd447 dlB.Scer\UAS.T:Ivir\HA1 medulla, with Scer\GAL4lz-gal4 dlGSd447 mesoderm dl15 mesothoracic tergum dl4 mesothoracic tergum | cold sensitive (with dl1) dl4 mesothoracic tergum | cold sensitive (with dl4) dl1 mesothoracic tergum | temperature conditional, with Scer\GAL4sca-537.4 dlUY2278 mushroom body, with Scer\GAL47B dlUY2278 dlUY3046 mushroom body, with Scer\GAL4ey-OK107 dlUY2278 mushroom body & axon, with Scer\GAL47B dlUY2278 dlUY3046 mushroom body & axon, with Scer\GAL4ey-OK107 dlUY2278 plasmatocyte | supernumerary, with Scer\GAL4Cg.PA dlScer\UAS.cHa segmental nerve, with Scer\GAL4scrt-11-6 dlGSd447 ventral furrow dl1 visceral mesoderm dl1 wing, with Scer\GAL4bbg-C96 dld05894
Classical Alleles ( 58 )
For All Classical Alleles Show
Allele of dl Class Mutagen Stocks Known lesion dl1 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate 4 -- dlk10816 P-element activity 2 -- dl4 amorphic allele - genetic evidence ethyl methanesulfonate 1 -- dl5-HA-2371 1 -- dld05894 1 -- dle04169 1 -- dlKG06652 P-element activity 1 -- dlUY2278 P-element activity 1 Yes dl15 amorphic allele - genetic evidence, loss of function allele X ray 0 -- dl6 amorphic allele - genetic evidence ethyl methanesulfonate X ray 0 -- dl8 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate X ray 0 -- dl01313 P-element activity 0 Yes dl10 loss of function allele ethyl methanesulfonate 0 -- dl11 ethyl methanesulfonate 0 -- dl12 loss of function allele X ray 0 -- dl1310 0 -- dl13 X ray 0 Yes dl14 ethyl methanesulfonate 0 -- dl16 loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes dl17 loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes dl18 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes dl19 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes dl20 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 -- dl21 loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes dl22 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes dl23 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes dl24 antimorphic allele - genetic evidence ethyl methanesulfonate 0 -- dl25 antimorphic allele - genetic evidence ethyl methanesulfonate 0 -- dl26 antimorphic allele - genetic evidence ethyl methanesulfonate gamma ray 0 -- dl27 antimorphic allele - genetic evidence, loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate gamma ray 0 Yes dl2 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes dl2L-136-13 ethyl methanesulfonate 0 -- dl2L-164-16 ethyl methanesulfonate 0 -- dl2L-258-38 ethyl methanesulfonate 0 -- dl2L-291-17 ethyl methanesulfonate 0 -- dl2L-294-9 ethyl methanesulfonate 0 -- dl2L-317-23 ethyl methanesulfonate 0 -- dl2L-325-31 ethyl methanesulfonate 0 -- dl2L-327-18 ethyl methanesulfonate 0 -- dl38 P-element activity 0 -- dl3 loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes dl5-SZ-3122 0 -- dl5 loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate X ray 0 Yes dl7 ethyl methanesulfonate X ray 0 -- dl9 ethyl methanesulfonate 0 -- dlc05444 piggyBac transposase 0 -- dlC447 ethyl methanesulfonate 0 Yes dld05429 P-element activity 0 -- dld08511 P-element activity 0 -- dlD447 ethyl methanesulfonate 0 Yes dlGS5251 0 Yes dlGSd447 0 Yes dlLY23 P-element activity 0 -- dlMI03808 0 -- dlQ10 0 -- dlQ17 ethyl methanesulfonate 0 -- dlunspecified 0 -- dlUY3046 P-element activity 0 Yes
Alleles Carried on Transgenic Constructs ( 98 )
For All Alleles Carried on Transgenic Constructs Show
Allele of dl Class Mutagen Stocks Known lesion dlGD1238 in vitro construct - RNAi in vitro construct - regulatory fusion 2 Yes dlScer\UAS.cHa in vitro construct - regulatory fusion 2 Yes dlHMS00028 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes dlHMS00727 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes dlJF02825 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes dlKK107820 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes dl+mWT in vitro construct - minigene 0 Yes dl1-184ΔNLS.Hsp83.T:Ecol\lacZ in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes dl138-342.Hsp83.T:Ecol\lacZ in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dl22.1.T:Zzzz\His6 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dl222-342.Hsp83.T:Ecol\lacZ in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dl3.Act5C in vitro construct - deletion 0 Yes dl327C.Hsp83.T:Ecol\lacZ in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dl327ΔNLSC.Hsp83.T:Ecol\lacZ in vitro construct - deletion 0 Yes dl380.Act5C in vitro construct - regulatory fusion 0 Yes dl380.T:boss-NT1 in vitro construct - coding region fusion 0 Yes dl380.WRPW.Act5C.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dl4.Act5C in vitro construct - deletion 0 Yes dl5.Act5C in vitro construct - deletion 0 Yes dl561.Act5C in vitro construct - amino acid replacement in vitro construct - deletion 0 Yes dl6.Act5C in vitro construct - deletion 0 Yes dl670.Act5C in vitro construct 0 Yes dl672.Act5C in vitro construct 0 Yes dl6XS-A in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dl7.Act5C in vitro construct - deletion 0 Yes dl8.Act5C in vitro construct 0 Yes dl84-342.Hsp83.T:Ecol\lacZ in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlAct5C.PA in vitro construct - regulatory fusion 0 Yes dlAct5C.PR in vitro construct - regulatory fusion in vitro construct - minigene 0 Yes dlAct5C.PS in vitro construct - regulatory fusion 0 Yes dlAct5C.PT in vitro construct - regulatory fusion 0 Yes dlAct5C.T:Hsim\VP16,T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlAct5C.T:Mmus\Rela,T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlAct5C.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlAla312.bcd in vitro construct - amino acid replacement 0 Yes dlB.Act5C in vitro construct - regulatory fusion 0 Yes dlB.Hsp83.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlB.Scer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlbcd.PB in vitro construct - regulatory fusion 0 Yes dlCTD.Hsp83.T:Scer\GAL4,T:boss-NT1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlCTDΔeh1.Hsp83.T:Scer\GAL4,T:boss-NT1 in vitro construct - deletion 0 Yes dlD.Act5C in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dldsRNA.cBa in vitro construct - RNAi 0 Yes dldsRNA.cSa in vitro construct - RNAi 0 Yes dlGlu312.bcd in vitro construct - amino acid replacement 0 Yes dlhs.PS in vitro construct - regulatory fusion 0 Yes dlHsp83.PG in vitro construct - regulatory fusion 0 Yes dlHsp83.Spe in vitro construct - regulatory fusion 0 Yes dlHsp83.T:boss-NT1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlHsp83.T:Ecol\lacZ in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlHsp83.T:Scer\GAL4,T:boss-NT1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlM16.T:boss-NT1 in vitro construct - amino acid replacement 0 Yes dlM18.T:boss-NT1 in vitro construct - amino acid replacement 0 Yes dlM2.T:boss-NT1 in vitro construct - amino acid replacement 0 Yes dlM20.T:boss-NT1 in vitro construct - amino acid replacement 0 Yes dlM23.T:boss-NT1 in vitro construct - amino acid replacement 0 Yes dlM4prime.T:boss-NT1 in vitro construct - amino acid replacement 0 Yes dlM7prime.T:boss-NT1 in vitro construct - amino acid replacement 0 Yes dlM8.T:boss-NT1 in vitro construct - amino acid replacement 0 Yes dlmC1.Hsp83 in vitro construct - amino acid replacement in vitro construct - regulatory fusion 0 Yes dlMtnA.T:Avic\GFP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlMtnA.T:Zzzz\His6 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlN.Hsp83.T:Ecol\lacZ in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlNLS.bcd in vitro construct - amino acid replacement 0 Yes dlNΔNLS.Hsp83.T:Ecol\lacZ in vitro construct - deletion 0 Yes dlQ.Act5C in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlRHR.Hsp83.T:Ecol\lacZ in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes dlRRKS.Act5C in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlS103A in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlS213A in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlS234P in vitro construct 0 Yes dlS312A in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlS317A in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlS70A in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlS79A in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlScer\UAS.cCa in vitro construct - regulatory fusion 0 Yes dlScer\UAS.cMa in vitro construct - regulatory fusion 0 Yes dlScer\UAS.P\T.2x.T:Avic\GFP-EGFP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlScer\UAS.P\T.cRa in vitro construct - regulatory fusion 0 Yes dlScer\UAS.P\T.T:Hsim\VP16 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlScer\UAS.P\T.T:Mmus\Rela in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlScer\UAS.T:Avic\GFP-EGFP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlScer\UAS.T:Avic\GFP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlScer\UAS.T:SV5\V5 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlScer\UAS.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlT290V in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlT85A in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlT:Avic\GFP in vitro construct - coding region fusion 0 Yes dlT:boss-NT1 in vitro construct - coding region fusion 0 Yes dlWRPW.Act5C.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlWRPW.Scer\UAS.P\T in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes dlY324F in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes dlαTub84B.PM in vitro construct - regulatory fusion 0 Yes dlΔC380.Act5C in vitro construct - regulatory fusion 0 Yes dlΔC470.Act5C in vitro construct - regulatory fusion 0 Yes dlΔC590.Act5C in vitro construct - regulatory fusion 0 Yes dlΔNLS.Hsp83 in vitro construct - regulatory fusion in vitro construct - site directed mutagenesis in vitro construct - deletion 0 Yes dlΔNLS antimorphic allele - genetic evidence in vitro construct - deletion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2L)BSC294 (Christensen and Cook, 2007.5.8) Df(2L)Exel7068 (Ryder, 2004.4.26) Df(2L)H20 (Steward and Nusslein-Volhard, 1986, Schupbach and Wieschaus, 1989, Schupbach and Wieschaus, 1991) Df(2L)H68 (Steward and Nusslein-Volhard, 1986) Df(2L)J4 (Huang et al., 2005, Chiu et al., 2005, Matova and Anderson, 2006) Df(2L)J4b (Rutschmann et al., 2000) Df(2L)M36F18 (Steward and Nusslein-Volhard, 1986) Df(2L)qua1374 (Mahajan-Miklos and Cooley, 1994) Df(2L)T317 (Steward and Nusslein-Volhard, 1986) Df(2L)TW119 (Huang et al., 2005, Chiu et al., 2005, Steward et al., 1984, Steward and Nusslein-Volhard, 1986, Tearle and Nusslein-Volhard, 1987, Govind et al., 1993, Kirov et al., 1993, Gillespie and Wasserman, 1994, Belvin et al., 1995, Lemaitre et al., 1995, Gross et al., 1996, Matova and Anderson, 2006) Df(2L)TW137 (Steward and Nusslein-Volhard, 1986, Tearle and Nusslein-Volhard, 1987, Schupbach and Wieschaus, 1989, Schupbach and Wieschaus, 1991) In(2L)dlH (Steward et al., 1984, Kirov et al., 1993, Gross et al., 1996) In(2L)dlT (Steward et al., 1984, Lemaitre et al., 1995)
Not disrupted in	Df(2L)M36F-S5 (Steward and Nusslein-Volhard, 1986, Tearle and Nusslein-Volhard, 1987) Df(2L)TW201 (Steward and Nusslein-Volhard, 1986) Df(2L)TW202 (Steward and Nusslein-Volhard, 1986) Df(2L)TW203 (Steward and Nusslein-Volhard, 1986) Df(2L)TW330 (Steward and Nusslein-Volhard, 1986) Df(2L)VA18 (Steward and Nusslein-Volhard, 1986)
Duplicated in	Dp(2;2)M(2)m+ (Wright, 1995.12.1)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD1238} NA P{KK107820} NA P{TRiP.HMS00028} NA P{TRiP.HMS00727} NA P{TRiP.JF02825} NA P{UAS-dl.3xFLAG} NA P{UAS-dl.B-HA} NA P{UAS-dl.C} NA P{UAS-dl.EGFP} NA P{UAS-dl.GFP} NA P{UAS-dl.H} NA P{UAS-dl.V5} NA P{UASp-2xEGFP.DLC} NA P{UASp-dl.M} NA P{UASp-dl.p65} NA P{UASp-dl.R} NA P{UASp-dl.VP16} NA P{UASp-dl.WRPW} NA heat-shock construct P{1-184ΔNLS-LacZ} NA P{84-342-LacZ} NA P{138-342-LacZ} NA P{222-342-LacZ} NA P{327.C-term-LacZ} NA P{327ΔNLSC-term-LacZ} NA P{dl-lacZ} NA P{dlRHR-lacZ} NA P{DorsalΔNLS} NA P{Hsp70-dl.S} NA P{Hsp83-dl.B.T:HA1} NA P{Hsp83-dl.nt1} NA P{hsp83-dl.Spe} NA P{hsp83-dl} NA P{hsp83-dlmC1} NA P{Hsp83-GAL4-dl.CTD.nt1} NA P{Hsp83-GAL4-dl.CTDΔeh1.nt1} NA P{Hsp83-GAL4-dl.nt1} NA P{N-term-LacZ} NA P{N-termΔNLS-LacZ} NA reporter construct P{1-184ΔNLS-LacZ} No P{84-342-LacZ} No P{138-342-LacZ} No P{222-342-LacZ} No P{327.C-term-LacZ} No P{327ΔNLSC-term-LacZ} No P{dl-lacZ} No P{dlRHR-lacZ} No P{N-term-LacZ} No P{N-termΔNLS-LacZ} No characterization construct P{bcd-dl.Ala312} NA P{bcd-dl.B} NA P{bcd-dl.Glu312} NA P{bcd-dl.NLS} NA P{dl.22.1} NA P{dl.380.nt1} NA P{dl.M2.nt1} NA P{dl.M4prime.nt1} NA P{dl.M7prime.nt1} NA P{dl.M8.nt1} NA P{dl.M16.nt1} NA P{dl.M18.nt1} NA P{dl.M20.nt1} NA P{dl.M23.nt1} NA P{dl.nt1} NA P{dl6XS-A} NA P{dl+mWT} NA P{dlS70A} NA P{dlS79A} NA P{dlS103A} NA P{dlS213A} NA P{dlS234P} NA P{dlS312A} NA P{dlS317A} NA P{dlT85A} NA P{dlT290V} NA P{dlY324F} NA P{dlΔNLS} NA P{dl-GFP.DL} NA P{αTub84B-dl.M} NA
Insertions
	Type of insertions Name Expression data insertion of mobile activating element P{EP}EP1158 NA P{GSV1}dlGSd447 NA P{GSV1}EP-MEP-M NA P{GSV2}dlGS5251 NA P{Mae-UAS.6.11}dlUY2278 NA P{Mae-UAS.6.11}dlUY3046 NA P{XP}dld05429 NA P{XP}dld05894 NA P{XP}dld08511 NA insertion of enhancer trap P{lacW}dlk10816 No P{lacW}l(2)SH1797SH1797 No P{PZ}dl01313 No miscellaneous insertions Mi{MIC}dlMI03808 NA P{hsp83-dl}38 NA P{RS5}dl5-HA-2371 NA P{RS5}dl5-SZ-3122 NA P{ry11}dlLY23 NA P{SUPor-P}dlKG06652 NA PBac{PB}dlc05444 NA PBac{RB}dle04169 NA
Gene Ontology: Function, Process & Cellular Component ( 40 unique terms )
Terms Based on Experimental Evidence ( 19 terms )
Molecular Function
CV term Evidence References
high mobility group box 1 binding inferred from physical interaction with Dsp1 (Decoville et al., 2000) protein binding inferred from physical interaction with Dip3 AND inferred from physical interaction with lwr AND inferred from physical interaction with smt3 (Bhaskar et al., 2000) inferred from physical interaction with tamo (Minakhina et al., 2003) repressing transcription factor binding inferred from physical interaction with Dsp1 (Decoville et al., 2000) inferred from physical interaction with gro (Valentine et al., 1998) RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in positive regulation of transcription inferred from direct assay (Thisse et al., 1991) RNA polymerase II regulatory region sequence-specific DNA binding inferred from direct assay (Ip et al., 1991, Valentine et al., 1998, Benveniste et al., 1998) contributes_to sequence-specific distal enhancer binding RNA polymerase II transcription factor activity inferred from genetic interaction with twi (Bhaskar and Courey, 2002) sequence-specific DNA binding inferred from direct assay (Copley et al., 2007)
Biological Process
CV term Evidence References
immune response inferred from expression pattern (Lemaitre et al., 1995) maternal determination of anterior/posterior axis, embryo inferred from mutant phenotype (Nusslein-Volhard, 1979) melanization defense response inferred from mutant phenotype (Bettencourt et al., 2004) negative regulation of gene expression inferred from direct assay (Anderson et al., 2006) peripheral nervous system neuron development inferred from mutant phenotype (Ayyar et al., 2007) plasmatocyte differentiation inferred from mutant phenotype (Huang et al., 2005) positive regulation of transcription from RNA polymerase II promoter inferred from direct assay (Benveniste et al., 1998) regulation of alternative nuclear mRNA splicing, via spliceosome inferred from mutant phenotype (Park et al., 2004) regulation of hemocyte proliferation inferred from direct assay (Matova and Anderson, 2006)
Cellular Component
CV term Evidence References
cytoplasm inferred from direct assay (Wu and Anderson, 1998, Bhattacharya and Steward, 2002) neuromuscular junction inferred from direct assay (Bolatto et al., 2003) nucleus inferred from direct assay (Wu and Anderson, 1998, Bhattacharya and Steward, 2002, Park et al., 2003)
Terms Based on Predictions or Assertions ( 25 terms )
Molecular Function
CV term Evidence References
DNA binding non-traceable author statement (Swiss-Prot Project Members, 1990.4.1) morphogen activity non-traceable author statement (Swiss-Prot Project Members, 1990.4.1) sequence-specific DNA binding transcription factor activity inferred from sequence or structural similarity with mouse Rel (FlyBase, 1992-) non-traceable author statement (Lemaitre, 2000.12.20)
Biological Process
CV term Evidence References
anterior/posterior pattern specification traceable author statement (Wylie, 1999) defense response non-traceable author statement (Lemaitre, 2000.12.20) traceable author statement (Williams et al., 1997) dorsal/ventral axis specification traceable author statement (Meister et al., 2000, Lall and Patel, 2001, Janssens and Beyaert, 2002) dorsal/ventral pattern formation non-traceable author statement (Martin-Blanco, 2000) ectodermal cell fate specification traceable author statement (Martin-Blanco, 2000) ectoderm development non-traceable author statement (Stathopoulos and Levine, 2002) gastrulation non-traceable author statement (Leptin, 1999) gastrulation involving germ band extension traceable author statement (Stathopoulos and Levine, 2004) germ cell migration traceable author statement (Wylie, 1999) heart development non-traceable author statement (Cripps and Olson, 2002) immune response non-traceable author statement (Swiss-Prot Project Members, 1990.4.1) traceable author statement (Martin-Blanco, 2000) mesodermal cell fate specification traceable author statement (Martin-Blanco, 2000) mesoderm development non-traceable author statement (Stathopoulos and Levine, 2002) negative regulation of transcription, DNA-dependent non-traceable author statement (Stathopoulos and Levine, 2002) traceable author statement (Courey and Jia, 2001) negative regulation of transcription from RNA polymerase II promoter traceable author statement (Lall and Patel, 2001) positive regulation of transcription, DNA-dependent traceable author statement (Courey and Jia, 2001) positive regulation of transcription from RNA polymerase II promoter traceable author statement (Cripps and Olson, 2002) regulation of transcription, DNA-dependent inferred from sequence or structural similarity with mouse Rel (FlyBase, 1992-) non-traceable author statement (Stathopoulos and Levine, 2002) Toll signaling pathway non-traceable author statement (Lemaitre, 2000.12.20) traceable author statement (Kimbrell and Beutler, 2001, Silverman, 2003, Leclerc and Reichhart, 2004) ventral cord development non-traceable author statement (Stathopoulos and Levine, 2002)
Cellular Component
CV term Evidence References
cytoplasm non-traceable author statement (Swiss-Prot Project Members, 1990.4.1) traceable author statement (Manfruelli et al., 1999, Meister et al., 2000, Lall and Patel, 2001) nucleus inferred by curator from GO:0000381 (Park et al., 2004) inferred by curator from GO:0003700 (FlyBase, 1992-) non-traceable author statement (van Eeden and St. Johnston, 1999, Swiss-Prot Project Members, 1990.4.1) traceable author statement (Manfruelli et al., 1999, Meister et al., 2000, Lall and Patel, 2001)
Sequence Ontology: Class of Gene
	gene (Minidorff et al., 2007) 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 dl allele Gene References dl1 cact (Wakabayashi-Ito et al., 2001) Dif (Stein et al., 1998) fog (Morize et al., 1998) lwr (Huang et al., 2005) wntD (Gordon et al., 2005) dl4 wntD (Gordon et al., 2005) dl15 sog (Araujo and Bier, 2000) dlS234P gd (Drier et al., 2000) dlunspecified gd (Drier et al., 2000) Taf1 (Pham and Sauer, 2000) unspecified cact (Ashburner et al., 1990) fs(1)Yb (Johnson et al., 1995) lwr (Chiu et al., 2005) Npc1b (Bhattacharya and Steward, 2003) pbl (Gregory et al., 2007) RpII140 (Parkhurst and Ish-Horowicz, 1991)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 61043 DroID - A comprehensive database of gene and protein interactions. • FBgn0260632 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 35047
Orthologs
Genome-wide drosophilid orthologs	Dana\GF15103 Dere\GG21746 Dgri\GH10077 Dmoj\GI16541 Dper\GL18753 Dpse\GA19765 Dsec\GM17128 Dsim\dl Dvir\GJ16232 Dwil\GK10525 Dyak\dl
Curated drosophilid orthologs	Dpse\GA19765   Dsim\dl   Dyak\dl
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL006930-PA Aedes aegypti (mosquito) XP_001949498.1 Acyrthosiphon pisum (pea aphid) ENSBTAP00000049624 Bos taurus (bovine) ENSCAFP00000006862 Canis familiaris (dog) CPIJ011758 Culex pipens (mosquito) ENSCPOP00000016204 Cavia porcellus (guinea pig) ENSCSAVP00000008151 Ciona savignyi (sedentary tunicate) 220294 Capitella spl (polychaete worm) ENSECAP00000018439 Equus caballus (horse) ENSP00000221452 Homo sapiens (human) ENSMMUP00000014199 Macaca mulatta (rhesus monkey) ENSMUSP00000092355 Mus musculus (house mouse) XP_001603465.1 Nasonia vitripennis (wasp) PHUM010722-PA Pediculus humanus (human body louse) ENSPPYP00000003533 Pongo pygmaeus (orangutan) ENSPTRP00000020529 Pan troglodytes (chimpanzee) ENSRNOP00000044552 Rattus norvegicus (norway rat) ENSTNIP00000016350 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000021998 Takifugu rubripes (pufferfish Takifugu) ENSXETP00000004697 Xenopus tropicalis (frog) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB12133 NV11353 LH23593 LH23563 FBgn0067431 FBgn0212540 FBgn0203417 FBgn0067500 FBgn0172039 JGI_V11_52849 FBgn0079761 FBgn0156354 FBgn0113924 FBgn0092128 CPIJ002469 BGIBMGA010496 ACYPI39891 ACYPI005133 ACYPI003588 GB30211 GB19537 GB18032 REL1 REL1A PHUM534140 GLEAN_07697
Stocks & Reagents
Stocks Listed in FlyBase ( 20 )
Bloomington	9319 y1 w*; P{UAS-dl.H}2 27650 y1 v1; P{TRiP.JF02825}attP2 32934 y1 sc* v1; P{TRiP.HMS00727}attP2 34938 y1 sc* v1; P{TRiP.HMS00028}attP2 7350 y1 w67c23; P{Mae-UAS.6.11}dlUY2278 3236 dl1 cn1 sca1/CyO, l(2)DTS1001
Harvard	- P{XP}dld05894 - PBac{RB}dle04169
Kyoto	125601 w1118; P{RS5}dl5-HA-2371
VDRC	v105491 P{KK107820}VIE-260B v45996 w1118; P{GD1238}v45996 v45998 w1118; P{GD1238}v45998
	More stocks available...
Genomic Clones ( 2 )
	BACR07M13 BACR21E16 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 39 )
	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	LP16165 RE58537
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	AT18533 GH07114 LD04247 LD18620 LD47815 RE23867 RE52892 RE56945 RE59372 RE65253 RH06445 SD23224
Other clones	71611 74069 97201 97853 157313 Dmel_dig_007108_0590_1833 EK173123 EK219757 EK228546 EK260237 EN06304 FGA002E06 FGC030H07 FGM323E05 RP004012968 RP004029684 RP004130861 RP004215212 RP004218434 RP004244617 RP004482733 RP004487456 RP004502354 RP004517368 RP004542206
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0260632 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 35047
Antibody Information
	polyclonal (Gross et al., 1999, Rushlow et al., 1989, Roth et al., 1989)
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of	Source for identity of: dl CG6667 (Zhong, 1999.11)
Source for database merge of	Source for merge of: dl anon-EST:GressD7 (Ashburner, 2004.6.30) Source for merge of: dl GSd447 (Minidorff et al., 2007)
Additional comments
Other Comments
	Gene expression is increased in response to the presence of two copies of Scer\GAL4hs.PB. (Liu and Lehmann, 2008) Ventral signal dependent modification of cact and dl may be required for the graded nuclear import of dl. (Drier et al., 2000) dl can activate transcription from zen and twi promoters, and additional Dsp1 inhibits the zen activation and increases the twi activation. (Brickman et al., 1999) dl protein is subject to signal-dependent phosphorylation while associated with cact protein in the cytoplasm. Phosphorylation of dl protein is required for its nuclear import. (Drier et al., 1999) Dif and dl are functionally redundant in their ability to control Drs gene expression in larvae. (Manfruelli et al., 1999) A linear activation cascade spz-Tl-cact-dl/Dif leads to the induction of the Drs gene in larval fat body cells. (Manfruelli et al., 1999) An interplay of dl and twi proteins with Taf4 protein is required for the activation of mesoderm-determining gene expression in the embryo. (Pham et al., 1999) The h and dl gene products continue to function as repressors in the setting up of segmentation in the absence of CtBP. (Nibu et al., 1998) Targeting of either tub or pll product to the plasma membrane by myristylation is sufficient to activate the signal transduction pathway that leads to translocation of the dl product. Activated Tl induces a localized recruitment of tub and pll proteins to the plasma membrane. (Towb et al., 1998) Tl pathway is required for the nuclear import of dl in the immune response, but not required for the nuclear import of Dif. Cytoplasmic retention of both dl and Dif depends on cact protein. The two signalling pathways that target cact for degradation must discriminate between cact-dl and cact-Dif complexes. (Wu and Anderson, 1998) Transcript induced by MMS treatment of S1CII cells. (Akaboshi and Inoue, 1997) nej is necessary for dl-mediated activation of the twi promoter. (Akimaru et al., 1997) dl protein binds specifically to the tub, pll and cact proteins. (Edwards et al., 1997) The molecular evolution of the Rel/NF-κB and IκB proteins is studied in parallel. Phylogenetic analysis allows the structure of the putative ancestor genes to be defined and proposes and evolutionary model that clusters both families in a unique Rel/NF-κB/IκB superfamily. (Huguet et al., 1997) dl and twi proteins synergistically activate transcription in cell culture from a promoter containing binding sites for both factors. The Rel homology domain of the dl protein appears to be sufficient for the synergy. Protein-protein interaction assays show that dl and twi proteins bind to one another in vitro. (Shirokawa and Courey, 1997) tub, pll, cact and dl form a complex essential for signal transduction. (Yang and Steward, 1997) Yeast two hybrid assay demonstrates both tub and pll interact with dl. Interactions have also been confirmed in an in vitro binding assay. (Yang and Steward, 1997) cact forms a concentration gradient inversely correlated to the nuclear translocation gradient of dl. Genetic evidence indicates degradation of cact is required, but not sufficient, to translocate dl completely into the nucleus. (Bergmann et al., 1996) Expression of dl causes lethality and the formation of melanotic tumours. (Govind, 1996) Homodimerisation, nuclear targeting and interaction of dl with cact are mediated by the conserved Rel-homology domain of dl. (Govind et al., 1996) dl and Dif have distinct DNA-binding characteristics and the proteins can heterodimerise in vitro. Mutants carrying no copies of dl and a single copy of Dif retain their full capacity to express the Dpt and CecA1 genes in response to bacterial challenge. (Gross et al., 1996) The embryonic regulatory pathway, comprising the gene products between spz and cact (Tl, tub and pll) but not the genes acting upstream or downstream (ea and dl), is involved in the induction of the Drs gene in adults. (Lemaitre et al., 1996) Expression of pll enhances the transcriptional activity and nuclear localisation of dl. (Norris and Manley, 1996) In embryos there is a gradient of cact protein. In ventral regions cact is degraded allowing dl to translocate into the nuclei, in dorsal regions cact persists, retaining dl in the cytoplasm. (Reach et al., 1996) The dl product binds to multiple sites in the dpp second intron. (Bai et al., 1995) cact inhibits nuclear translocation of dl on the ventral side of the embryo by binding to and retaining dl in the cytoplasm. cact is rapidly degraded in response to signalling from the dorsal ventral pathway between spz and dl/cact, this signal-dependent degradation does not require the presence of dl but does require sequences in the amino terminus or ankyrin repeats of the cact protein. Disruption of the dl-cact complex is a secondary result of cact degradation. (Belvin et al., 1995) Monitoring phosphorylation in mutant backgrounds demonstrates that phosphorylation of the dl protein is clearly affected. Therefore phosphorylation may play a role in regulating the dl protein. One of three mutations in the putative phosphorylation sites at residue 312, not at residue 290 or 324, markedly reduces the ability of the transgene to rescue dl mutant embryos. (Drier and Steward, 1995) Molecular analysis suggests that sna protein acts over distances of 50-150bp to block the activity, but not the binding, of the dl activator to the rho 650bp enhancer. (Gray et al., 1995) dl protein enhances the biosynthesis and stability of cact. (Kubota and Gay, 1995) Increases in intra-cellular Ca2+ levels result in rapid destruction of cact protein and dephosphorylation of dl protein in a Drosophila cell line. (Kubota and Gay, 1995) cact activity blocks the DNA binding and nuclear localisation functions of dl. dl transcriptional activating region is functional in the dl cact complex. (Lehming et al., 1995) Nuclear localisation of dl in the fat body during the immune response is controlled by the intracellular embryonic dorsoventral pathway, the Tl signalling pathway. dl alone does not control the expression of antibacterial peptide genes as these genes are inducible in its absence. dl is not involved in the formation of melanotic tumours of Tl or cact mutations nor in the induction of Dpt gene, or dl acts in concert with other proteins to affect cellular and/or humoral immunity. (Lemaitre et al., 1995) The dl regulatory gradient defines the limits of inductive interactions between germ layers after gastrulation. (Maggert et al., 1995) The tub protein can function in a novel way to enhance dl activity. In the absence of dl or when dl is cytoplasmic, tub is only found in the cytoplasm of transfected cells. When dl is localised to the nucleus, so is tub. tub can then function to enhance reporter gene expression, by cooperation with dl or as a Scer\GAL4-tub fusion protein. tub is capable of acting as both a chaperon or escort for dl as it moves to the nucleus and then as a transcriptional coactivator. The intracytoplasmic domain of Tl is sufficient for activating the signalling pathway that leads to dl-tub nuclear translocation in Schneider cells. (Norris and Manley, 1995) dl can activate the CecA1 promoter, but to a lesser extent and in a less sequence-specific manner than Dif. The dl product exerts a dominant negative effect on Dif transactivation of CecA1. (Petersen et al., 1995) The dl-bHLH protein interactions mediating gene expression in the neuroectoderm and mesoderm are fundamentally distinct. Proximity between dl and bHLH binding sites is essential for synergistic activation of gene expression in the lateral neuroectoderm, where levels of dl product diminish. Sharp on/off patterns of gene expression in the presumptive mesoderm do not require linkage of these sites. (Szymanski and Levine, 1995) A sequence within the RHD (Rel homology domain) is essential for inhibitor interactions. Point mutations within this sequence can uncouple DNA binding and inhibitor interactions in vitro. (Tatei and Levine, 1995) dl is an embryonic phosphoprotein and its phosphorylation state is regulated by an intracellular signaling pathway initiated by the transmembrane receptor Tl. Using a combined genetic and biochemical approach it is demonstrated that activation of Tl stimulates an increase in the extent of dl phosphorylation. Signal-dependent dl phosphorylation is modulated by three intracellular proteins, pll, tub and cact. (Gillespie and Wasserman, 1994) Dorsal-ventral patterning is regulated by a signalling pathway that includes Tl and transcription factors, dl, that interact with related enhancers, rho. The κ enhancer from mouse is capable of generating lateral stripes of Ecol\lacZ gene expression in transgenic embryos in a pattern similar to that directed by rho enhancer. Results suggest that enhancers can couple conserved signalling pathways to divergent gene functions, dorso-ventral patterning and mammalian haematopoiesis. (Gonzalez-Crespo and Levine, 1994) ems, a head-specific gap gene, may function as a co-repressor of dl, thereby linking the anteroposterior and dorsoventral systems st the molecular level. (Grossniklaus et al., 1994) Disulfide cross-linking in crude extracts has identified two complexes of dl protein: a dl protein homodimer and a complex of the homodimer with cact protein. The distribution of the complex varies, the homodimer is the nuclear form of dl protein and a complex of the homodimer with cact protein prevails in the cytoplasm. (Isoda and Nusslein-Volhard, 1994) dl directly represses tld gene expression in ventral regions of the early embryo. DNaseI footprinting reveals dl binds to at least three sites in the tld 5' flanking sequences that are required in vivo for ventral repression (423bp VRE element). (Kirov 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 Tl signalling pathway generates a dl nuclear gradient which initiates the differentiation of the mesoderm, neuroectoderm and dorsal ectoderm by activating and repressing gene expression in the early embryo. A second signalling pathway controlled by the tor receptor kinase also modulates dl activity. The tor pathway selectively masks the ability of dl to repress gene expression but only has a slight effect on activation. (Rusch and Levine, 1994) dl gene product interacts with members of the HLH family, including da, ac and sc and dosage sensitive interactions that exist between dl, da, ac, sc and twi are required for the specification of both the embryonic mesoderm and neuroectoderm. (Gonzalez-Crespo and Levine, 1993) Promoter fusions using elements of the twi, ve, da and sna promoters indicate that low affinity dl-binding sites restrict target gene expression to the presumptive mesoderm, where there are peak levels of dl expression, while high affinity sites in other target genes permit expression in ventrolateral regions where dl levels are intermediate. Activation by low levels of dl in lateral regions depends on cooperative interaction between dl and other basic helix loop helix proteins. Promoters containing the Et (veinlet) or Eds (dl and snail) E boxes display opposite behaviour in da and twi mutants, suggesting they are regulated by different basic helix loop helix proteins. (Jiang and Levine, 1993) A minimal 110bp Ventral Repression Element silencer in the zen promoter contains two dl binding sites as well as binding sites for additional nuclear factors present in early embryos. Mutations in the latter convert the minimal VRE into an enhancer, mediating transcriptional activation in ventral regions in response to dl. Thus dl is converted from an activator to a silencer by interactions with neighboring corepressors. (Jiang et al., 1993) dl binding sites from the zen promoter can mediate transcriptional activation of a heterologous promoter, but not repression. T-rich sequences close to the dl binding sites in the silencer region of the zen promoter are conserved between D.melanogaster, D.virilis and D.pseudoobscura.pseudoobscura. (Kirov et al., 1993) Increased cytoplasmic calcium concentration and the expression of constitutively active Tl receptors can induce the relocalisation of dl in culture cells. Activation of endogenous Pka-C1, expression of wild type Tl receptors or treatment of cells with activators of Pkc53E and radical oxygen intermediates have only a marginal effects on the cellular distribution of dl protein. (Kubota et al., 1993) Ecol\lacZ reporter gene constructs demonstrate the presence of dl maternal system cis-acting response elements in the 5' flanking region of tll. (Liaw and Lengyel, 1993) In addition to its role in embryogenesis, dl is involved in the immune response. In dl mutants the genes encoding antibacterial peptides retain their inducibility, suggesting multifactorial control. (Reichhart et al., 1993) Cytoplasmic injection studies indicate that the putative Toll ligand appears to originate from a ventrally restricted zone extending along the anterior-posterior axis, and its diffusion or graded release are required to determine the slope of the nuclear dorsal protein gradient. (Roth, 1993) dl and cact are phosphoproteins that form a stable cytoplasmic complex. The cact protein is stabilised by its interaction with dl protein, and the dl-cact complex dissociates when dl protein is targeted to the nucleus. (Whalen and Steward, 1993) The zygotic dpp gradient and the maternal dorsal gradient specify distinct, non-overlapping domains of the dorsal-ventral pattern. (Wharton et al., 1993) Double mutant combinations of dl with ea alleles demonstrate that spatial regulation of ea activity by localized zymogen activation is a key initial event in defining the polarity of the dorsal-ventral embryonic pattern. (Chasan et al., 1992) In vitro studies showed the cactus gene product can inhibit binding of dorsal protein to DNA. (Geisler et al., 1992) Expression of dl-lacZ fusion protein causes a partial loss of function dl phenotype, and downstream genes twi and zen are misregulated. The fusion protein localises to nuclei in manner indistinguishable from wild type dl protein. The dl-lacZ fusion provides some dl function: in a null dl background the fusion gene causes partial rescue of the phenotype. In vitro immunoprecipitation experiments show that dl and dl-lacZ proteins associate, suggesting that in wild type dl functions as an oligomer. Dominant female sterility caused by dl-lacZ is relieved by additional dl or Ecol\lacZ protein. (Govind et al., 1992) dl acts in concert with basic HLH proteins (possibly including twi) to activate ve in both ventral and lateral regions. A dl activator site has been found in the neural ectoderm expression region of the ve promoter. (Ip et al., 1992) Mutants of dl have been sequenced and their phenotypes studied. Results demonstrate that the dl protein has an amino terminal DNA binding domain and a carboxy terminal domain required for transcriptional activation/repression. (Isoda et al., 1992) dl binding site domain exchange experiments, using Ecol\lacZ reporter gene constructs, between the zen and twi promoters demonstrate that dl is intrinsically an activator and that repression requires additional factors present in the distal region of the zen promoter, the VR. (Jiang et al., 1992) In vitro mutagenesis alleles assayed in a cell free system indicate that the rel homology domain of dl, around amino acid 270, is the site at which the cact gene product binds. (Kidd, 1992) Effects of Toll on dl in cotransfected Schneider cells examined: Toll can enhance nuclear localisation of dl and, independently, the ability of dl to activate transcription once in the nucleus. Pathway from Toll to dl may involve protein kinase A, and nuclear transport and activation of dl may result from phosphorylation of dl by protein kinase A. (Norris and Manley, 1992) The dl gene product is required for ventral repression of tll expression of the stripe: bcd function is also required. (Pignoni et al., 1992) The dl gene acts downstream of Tl: perivitelline fluid from dl mutant embryos is equivalent to that obtained from wild type embryos. (Stein and Nusslein-Volhard, 1992) The effect of the terminal system on the expression of 2 zygotic genes involved in dorsoventral patterning, sna and dpp, is mediated by a reduction in dl activity by the terminal system. Due to this interaction the poles adopt a more dorsalised fate than their counterparts in the middle of the embryo. (Casanova, 1991) dl is a sequence specific DNA binding protein that may mediate long range repression by interacting with the distal regions of the zen promoter. dl binding sites share sequence similarity with the conserved sites recognized by the rel and NF-ΚB proteins. (Ip et al., 1991) A combination of promoter fusion-P-element transformation assays (1.2kb twi promoter fragment is sufficient to generate normal twi pattern of Ecol\lacZ expression) and in vitro DNA binding assays coupled with site directed mutagenesis (revealing four dl-binding sites in the twi promoter) have been used to establish a link between the dl-binding sites and twi expression in early embryos. The dorsal ventral limits of twi expression depend on the number and affinity of dl binding sites present in the twi promoter. dl-binding sites present in the twi promoter possess a lower affinity to those present in the zen promoter. (Jiang et al., 1991) Establishment of the mesoderm neuroectoderm boundary involves the interaction of twi, sna and dl proteins. (Kosman et al., 1991) Footprint analysis has been used to analyse transcription activation factors responsible for ventral specific expression of twi. dl has been found to bind the ventral activation region of twi and interact directly or indirectly with other DNA bound regulatory factors to activate twi expression in the presumptive mesoderm. (Pan et al., 1991) Mutations in maternal dorsal class gene dl interact with RpII140wimp. (Parkhurst and Ish-Horowicz, 1991) dl appears to activate the expression of twi and sna and repress the expression of zen and dpp. Polar expression of dpp and zen requires the terminal system to override the repression of dl, and twi and sna polar expression require the terminal system to augment activation of dl. (Ray et al., 1991) dl acts as a sequence specific trans-activator of the twi promoter. Gel retardation assays have been used to investigate binding of dl protein to synthetic oligonucleotides corresponding to the proximal and distal activator region of the twi promoter. (Thisse et al., 1991) Involved in the regulatory hierarchy responsible for the asymmetric distribution and function of zygotic regulatory gene products along the DV axis of early embryos. (Rushlow and Levine, 1990) Establishment of the dl gradient involves selective nuclear transport. Truncated dl protein lacking C-terminal sequences accumulate predominantly in the nuclei of transfected Schneider cells while the full length protein is largely restricted to the cytoplasm. (Rushlow et al., 1989) Mutations in dl result in a maternal effect phenotype with defects during the early stages of gastrulation and defects in the dorsoventral axis; embryos derived from homozygous females are dorsalised. (Schupbach and Wieschaus, 1989) Epistatic relationships exist between dorsalizing maternal effect mutations and "dppHin" alleles. (Irish and Gelbart, 1987) opa-e is present at position 1417bp, identified as a stretch of 34 glutamine residues, within the dl locus. (Steward, 1987) The expression of genes controlling neurogenesis is dependent on the previous activity of the genes controlling the development of the embryonic dorsal-ventral pattern. (Campos-Ortega, 1983) In homozygous embryos invagination of the ventral presumptive mesodermal cells fails to occur and the resulting embryos are devoid of internal organs. (Simpson, 1983) Embryos produced by homozygous dl females form normal cellular blastoderm but at gastrulation develop into yolk-filled tube of dorsal hypoderm. Hair pattern of cuticle characteristic of dorsal hypoderm; ventral structures, such as denticle belts, lacking. Normally, dorsal infoldings occupy entire circumference of embryo. Evidence of anterior-posterior differentiation includes possible mouth armature structures anteriorly, small spiracles posteriorly and orientation of hairs. The periodicity of stripes of ftz expression in pre-gastrulation embryos, as revealed by antibody staining, displays the pattern normally characteristic of the dorsum circumferentially in embryos produced by dl females (Carroll, Winslow, Twombly, and Scott, 1987). Embryos produced by dl/dl and dl/Df(2L)TW137 indistinguishable, suggesting dl to be amorphic. Penetrance complete; expression constant. Embryos of dl2 females lack all structures normally derived from the ventral half of the egg, including mesoderm, endodermal gut, ventral nervous system and ventral hypoderm. dl1 and to a lesser extent dl2, females produce embryos with reduced capacity for neurogenesis in response to an absence of dl function (Campos-Ortega, 1983). dl germ line-dependent; homozygous germ-line clones produce dorsalized embryos (Schupbach and Wieschaus, 1986). Embryos of dl/+ females produced at 29oC develop into comparatively normal-looking larvae; they mainly lack internal organs, such as mesoderm and parts of the anterior and posterior gut; often ventral hypoderm including denticle belts reduced; phenotype sensitive to genetic background. At 22oC, dl/+ females produce normal embryos. Developmental fate of ventrally located cells on cellular blastoderm apparently shifted to that of more dorsally located cells. The phenotype of embryos produced by dl/dl females partially rescued by the injection of wild-type cytoplasm but not RNA (Santamaria and Nusslein-Volhard, 1983; Anderson and Nusslein-Volhard, 1984). Developmental profiles show transcript to be present only in ovaries and pre-cellular-blastoderm stages of embryogenesis. In situ hybridization indicates that ovarian transcript accumulates in nurse cells from stage 5 to 11; number of transcripts per genome equivalent in these polytene cells remains low and constant until stage 10, at which time there is a dramatic increase in the relative numbers of transcripts. After a lag of one or two nuclear divisions, transcript begins to accumulate in the oocyte; by stage 12 there is little detectable transcript in the nurse cells. It appears as though the nurse-cell transcript is transferred to the oocyte and thus to the embryo; transcript seems to be uniformly distributed in stage 14 oocytes (Steward, Ambrosio, and Schedl, 1985). dl protein is uniformly distributed throughout cytoplasm of early embryo; in the syncytial blastoderm a gradient of expression is achieved by the graded transport of dl protein into nuclei, with the highest nuclear concentrations found ventrally; protein remains cytoplasmic dorsally. Maternal dorsalizing mutants prevent nuclear localization and ventralized embryos show dorsal as well as ventral nuclear localization (Steward, Zusman, Huang and Schedl, 1988; Rushlow, Han, Manley and Levine, 1989; Steward, 1989; Roth et al., 1989).
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_165217 NM_165218 NM_165219 NM_001169529 NM_001169530 RefSeq (Proteins) • NP_724052 NP_724053 NP_724054 NP_001163000 NP_001163001 DDBJ / EMBL / GenBank • AAA28479 AAC35296 AAF53610 AAF53611 AAF53612 AAT94434 ACZ94286 ACZ94287 AF053614 AQ025943 BH900897 BI372084 BT015205 CZ469985 CZ474527 CZ476269 M23702 X65272 Entrez Gene - A searchable database of RefSeq genes. • 35047 UniProtKB/Swiss-Prot • P15330
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • NF-kappa-B/Rel/dorsal (IPR000451) Cell surface receptor IPT/TIG (IPR002909) p53-like transcription factor, DNA-binding (IPR008967) Rel homology (IPR011539) Immunoglobulin-like fold (IPR013783) Immunoglobulin E-set (IPR014756)
Linkouts
	BioGRID - A database of protein and genetic interactions • 61043 DroID - A comprehensive database of gene and protein interactions. • FBgn0260632 DRSC - Results from RNAi screens. • FBgn0260632 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0260632 FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0260632 FlyReactome - A curated knowledgebase of Drosophila melanogaster pathways • 89278 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 35047 InParanoid A subset of ortholog calls from InParanoid. • AAEL006930-PA Aedes aegypti (mosquito) XP_001949498.1 Acyrthosiphon pisum (pea aphid) ENSBTAP00000049624 Bos taurus (bovine) ENSCAFP00000006862 Canis familiaris (dog) CPIJ011758 Culex pipens (mosquito) ENSCPOP00000016204 Cavia porcellus (guinea pig) ENSCSAVP00000008151 Ciona savignyi (sedentary tunicate) 220294 Capitella spl (polychaete worm) ENSECAP00000018439 Equus caballus (horse) ENSP00000221452 Homo sapiens (human) ENSMMUP00000014199 Macaca mulatta (rhesus monkey) ENSMUSP00000092355 Mus musculus (house mouse) XP_001603465.1 Nasonia vitripennis (wasp) PHUM010722-PA Pediculus humanus (human body louse) ENSPPYP00000003533 Pongo pygmaeus (orangutan) ENSPTRP00000020529 Pan troglodytes (chimpanzee) ENSRNOP00000044552 Rattus norvegicus (norway rat) ENSTNIP00000016350 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000021998 Takifugu rubripes (pufferfish Takifugu) ENSXETP00000004697 Xenopus tropicalis (frog) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /torstoll/dorsal1.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 35047 modMine - Data generated by the modENCODE project. • FBgn0260632 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB12133 NV11353 LH23593 LH23563 FBgn0067431 FBgn0212540 FBgn0203417 FBgn0067500 FBgn0172039 JGI_V11_52849 FBgn0079761 FBgn0156354 FBgn0113924 FBgn0092128 CPIJ002469 BGIBMGA010496 ACYPI39891 ACYPI005133 ACYPI003588 GB30211 GB19537 GB18032 REL1 REL1A PHUM534140 GLEAN_07697 REDfly - A database of transcriptional regulatory elements. • FBgn0260632
Synonyms & Secondary IDs ( 14 )
Reported As
Symbol Synonym	anon-EST:GressD7   CG6667 (Ashburner, 2004.6.30, De Gregorio et al., 2001, Molnar et al., 2006, Carrera et al., 2008, Keleman et al., 2009.8.5, Perkins et al., 2009, Hernandez-Romano et al., 2008, Ni et al., 2010.12.1, Ni et al., 2011.7.19, Cammarato et al., 2011) d; (Crocker et al., 2008) dl (Bhaskar et al., 2002, Lehmann et al., 2002, Aerts et al., 2007, Cantera et al., 1999, Ayyar et al., 2007, Matova and Anderson, 2010, Zeitouni et al., 2007, Minakhina and Steward, 2006, Carneiro et al., 2006, Xavier-Neto, 2005, Scuderi et al., 2006, Christensen and Cook, 2007.5.8, Anderson et al., 2006, Ratnaparkhi et al., 2006, Uvell and Engstrom, 2003, Mizutani et al., 2006, Matova and Anderson, 2006, Gim et al., 2001, Kalamarz et al., 2008, Molnar et al., 2006, Liu and Lehmann, 2008, Prothmann et al., 2006, Minidorff et al., 2007, DeLotto et al., 2007, Bornemann et al., 2008, Pope and Harris, 2008, Heckscher et al., 2007, Tan et al., 2008, Liberman and Stathopoulos, 2009, Carrera et al., 2008, Dworkin et al., 2009, Boettiger and Levine, 2009, Luo et al., 2001, Beramendi et al., 2007, Thoetkiattikul et al., 2005, Shirangi et al., 2006, Liberman et al., 2009, Fontenele et al., 2009, Bhattacharya and Steward, 2002, Obbard et al., 2009, Steneberg and Samakovlis, 2001, Yano et al., 2008, Salzer et al., 2010, Blanco and Gehring, 2008, Mrinal and Nagaraju, 2010, Jia et al., 2002, Paddibhatla et al., 2010, Aerts et al., 2010, van Uden et al., 2011, Ajuria et al., 2011, The modENCODE Consortium, 2010, The modENCODE Consortium, 2010, Negreiros et al., 2010) Dl (Sample and Shvartsman, 2010, Chen, 2006, Biemar et al., 2006, Park et al., 2003, Kanodia et al., 2009, Xi et al., 2008, Fakhouri et al., 2010) dL (Capy et al., 2000) DL (Ratnaparkhi and Courey, 2008, MacArthur et al., 2009) Dor (Rand et al., 2010) fs(2)k10816 (Spradling, 1999.12.9) GSd447   mat(2)dorsal
Name Synonym	Dorsal (Bhaskar et al., 2002, Lehmann et al., 2002, Matova and Anderson, 2010, Sample and Shvartsman, 2010, Sivatchenko and Letsou, 2006, Nibu et al., 2007, McElwain et al., 2007, Lawrence, 2006, Akira et al., 2006, Chen et al., 2006, Chen, 2006, Zinzen et al., 2006, Biemar et al., 2006, Hogarth et al., 2005, Nguyen and Frasch, 2006, Bergmann, 2006, Zeitlinger et al., 2007, Minakhina et al., 2003, Chanut, 2006, Stathopoulos and Levine, 2005, Uvell and Engstrom, 2003, Waterhouse et al., 2007, Rutschmann et al., 2000, Matova and Anderson, 2006, Chen et al., 2000, Avila et al., 2002, Park et al., 2003, Kuranaga and Miura, 2007, Harari-Steinberg et al., 2007, Araujo et al., 2008, Levine et al., 2008, Ratnaparkhi and Courey, 2008, Nibu et al., 2003, Cho, 2004, Roth et al., 2003, Christophides et al., 2002, Cowden and Levine, 2003, Ching et al., 2008, DeLotto et al., 2007, Reed et al., 2008, Qian et al., 2011, Tan et al., 2008, Xylourgidis et al., 2006, Mavrakis et al., 2009, Korolchuk et al., 2007, Liang et al., 2008, Buchon et al., 2009, Jia et al., 2002, Nie et al., 2009, Towb et al., 2009, Beramendi et al., 2007, Cronin et al., 2009, MacArthur et al., 2009, Kanodia et al., 2009, Fontenele et al., 2009, Copley et al., 2007, Khoueiry et al., 2010, Xi et al., 2008, Tanji et al., 2010, Kechris and Li, 2008, Tipping et al., 2010, Paddibhatla et al., 2010, van Uden et al., 2011, Wang et al., 2011, Fakhouri et al., 2010, Rand et al., 2010) dorsal (Cantera et al., 1999, Ayyar et al., 2007, Kankel et al., 2007, Zeitouni et al., 2007, Tanji et al., 2007, Matova and Anderson, 2007, Shen and Tanda, 2007, Fu and Levine, 2007, Boulay et al., 1987, Gregory et al., 2007, Wu and Silverman, 2007, Senger et al., 2006, Anderson et al., 2006, Gim et al., 2001, Fu and Levine, 2008, Meinhardt, 2004, Bolatto et al., 2003, Prothmann et al., 2006, Busse et al., 2007, Bosveld et al., 2008, Crocker et al., 2008, Yajima et al., 2003, Diangelo et al., 2009, Levine et al., 2007, Luo et al., 2001, Schaaf et al., 2009, Shirangi et al., 2006, Liberman et al., 2009, Bhattacharya and Steward, 2002, Steneberg and Samakovlis, 2001, Yano et al., 2008, Takano and Gusella, 2002, Beckstead et al., 2005, Salzer et al., 2010, Goto et al., 2010, Blanco and Gehring, 2008, Mosca and Schwarz, 2010, Mrinal and Nagaraju, 2010, Valanne et al., 2010, Bettencourt et al., 2004, Zsindely et al., 2009, Beramendi et al., 2005, Aerts et al., 2010, Ajuria et al., 2011, Cornwell and Kirkpatrick, 2001, Cammarato et al., 2011) Embryonic polarity protein dorsal (Hernandez-Romano et al., 2008)
Secondary FlyBase IDs
	FBgn0000462 FBgn0025325 FBgn0260285
References ( 708 )
Generate a list of	All references relating to this gene ( 708 )
List References by type	Research paper  ( 359 ) Supplementary material  ( 10 ) Review  ( 196 ) Personal communication to FlyBase  ( 13 ) Abstract  ( 109 ) FlyBase analysis  ( 2 ) Conference report  ( 7 ) Stock list  ( 1 ) Curated genome annotation  ( 2 ) Computer file  ( 4 ) Protein sequence record  ( 1 ) Automatic genome annotation  ( 1 ) Book  ( 1 ) Biography  ( 1 )
Recent research papers ( 21 )
	Ajuria et al., 2011, Development 138(5): 915--924 Capicua DNA-binding sites are general response elements for RTK signaling in Drosophila. [FBrf0212975] Qian et al., 2011, Genomics 97(5): 294--303 The full-length transcripts and promoter analysis of intergenic microRNAs in Drosophila melanogaster. [FBrf0213652] Sayal et al., 2011, Fly 5(1): 47--52 Optimization of reporter gene architecture for quantitative measurements of gene expression in the Drosophila embryo. [FBrf0213005] van Uden et al., 2011, PLoS Genet. 7(1): e1001285 Evolutionary Conserved Regulation of HIF-1β by NF-κB. [FBrf0212948] Wang et al., 2011, Infect. Immun. 79(2): 606--616 Host and Pathogen Glycosaminoglycan-Binding Proteins Modulate Antimicrobial Peptide Responses in Drosophila melanogaster. [FBrf0212806] Aerts et al., 2010, PLoS Biol. 8(7): e1000435 Robust Target Gene Discovery through Transcriptome Perturbations and Genome-Wide Enhancer Predictions in Drosophila Uncovers a Regulatory Basis for Sensory Specification. [FBrf0211419] Fakhouri et al., 2010, Mol. Syst. Biol. 6: 341 Deciphering a transcriptional regulatory code: modeling short-range repression in the Drosophila embryo. [FBrf0209745] Goto et al., 2010, J. Biol. Chem. 285(21): 15731--15738 Cooperative regulation of the induction of the novel antibacterial Listericin by peptidoglycan recognition protein LE and the JAK-STAT pathway. [FBrf0210770] Khoueiry et al., 2010, Curr. Biol. 20(9): 792--802 A cis-regulatory signature in ascidians and flies, independent of transcription factor binding sites. [FBrf0210780] Matova and Anderson, 2010, J. Cell Sci. 123(4): 627--633 Drosophila Rel proteins are central regulators of a robust, multi-organ immune network. [FBrf0209910] Mosca and Schwarz, 2010, Nat. Neurosci. 13(8): 935--943 The nuclear import of Frizzled2-C by Importins-beta11 and alpha2 promotes postsynaptic development. [FBrf0211361] Mrinal and Nagaraju, 2010, J. Biol. Chem. 285(31): 24206--24216 Dynamic Repositioning of Dorsal to Two Different {kappa}B Motifs Controls Its Autoregulation during Immune Response in Drosophila. [FBrf0211351] Negreiros et al., 2010, genesis 48(1): 31--43 alphaPS1betaPS integrin receptors regulate the differential distribution of Sog fragments in polarized epithelia. [FBrf0209674] Paddibhatla et al., 2010, PLoS Pathog. 6(12): e1001234 Role for sumoylation in systemic inflammation and immune homeostasis in Drosophila larvae. [FBrf0212643] Perry et al., 2010, Curr. Biol. 20(17): 1562--1567 Shadow enhancers foster robustness of Drosophila gastrulation. [FBrf0211793] Rand et al., 2010, Insect Biochem. Mol. Biol. 40(11): 792--804 Permeabilization of Drosophila embryos for introduction of small molecules. [FBrf0211955] Salzer et al., 2010, Genetics 184(1): 185--197 The retinal determination gene eyes absent is regulated by the EGF receptor pathway throughout development in Drosophila. [FBrf0209690] Sample and Shvartsman, 2010, Proc. Natl. Acad. Sci. U.S.A. 107(22): 10092--10096 Multiscale modeling of diffusion in the early Drosophila embryo. [FBrf0211013] Tanji et al., 2010, Proc. Natl. Acad. Sci. U.S.A. 107(33): 14715--14720 Heterodimers of NF-{kappa}B transcription factors DIF and Relish regulate antimicrobial peptide genes in Drosophila. [FBrf0211616] Tipping et al., 2010, EMBO J. 29(19): 3222--3235 β-arrestin Kurtz inhibits MAPK and Toll signalling in Drosophila development. [FBrf0211995] Valanne et al., 2010, J. Immunol. 184(11): 6188--6198 Genome-wide RNA interference in Drosophila cells identifies G protein-coupled receptor kinase 2 as a conserved regulator of NF-kappaB signaling. [FBrf0210846] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...