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Gene Dmel\tor

General Information
Symbol	Dmel\tor	Species	D. melanogaster
Name	torso	Annotation symbol	CG1389
Feature type	protein_coding_gene	FlyBase ID	FBgn0003733
Gene Model Status	Current	Stock availability	25 publicly available
Also Known As	Torso
Genomic Location
Chromosome (arm)	2R	Recombination map
Cytogenetic map	43E11-43E12	Sequence location	2R:3,590,680..3,597,042 [-]
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 torso is referred to in FlyBase by the symbol Dmel\tor (CG1389, FBgn0003733). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: protein tyrosine kinase activity. There is experimental evidence that it is involved in the biological process: terminal region determination; protein autophosphorylation; pole cell migration; metamorphosis; gastrulation; chorion-containing eggshell pattern formation. 137 alleles are reported. The phenotypes of these alleles are annotated with: organ system subdivision; organ system; multicellular structure; primordium; larval head segment; portion of tissue; embryonic segment; abdominal segment 7; mouthpart; epithelial furrow; cephalopharyngeal skeleton. It has 3 annotated transcripts and 3 annotated polypeptides. Protein features are: Fibronectin, type III; Protein kinase, ATP binding site; Protein kinase, catalytic domain; Protein kinase-like domain; Serine-threonine/tyrosine-protein kinase; Tyrosine-protein kinase, active site; Tyrosine-protein kinase, catalytic domain; Tyrosine-protein kinase, receptor torso. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of extremely low expression. Peak expression observed within 00-06 hour embryonic stages, in adult female stages. Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: adult ovary. Comments on Affy2 ProbeSet: ProbeSet 1638587_at completely aligns to an exonic region common to each of the 3 FlyBase-annotated transcript isoforms of tor. Gene sequence location is 2R:3590680..3597042.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	tor: torso Maternal-effect lethal; embryos from homozygous mothers show alterations in the anterior-posterior pattern. Hypoactivity (loss-of-function) mutant embryos lack anterior-most head structures (labrum, dorsal bridges) as well as structures posterior to the seventh abdominal segment. Hyperactivity (gain of function) mutant embryos, on the other hand, show segment defects in the middle of the embryos, but may have enlarged terminal structures (Klingler et al., 1988; Strecker et al., 1989). A large number of revertants have been obtained from dominant or semi-dominant hypermorphic alleles. During cellularization at the blastoderm stage, hypoactivity mutant embryos show a "pole hole" phenotype. A funnel of yolk-free cytoplasm with a small number of nuclei (between 10 and 20) is formed at the posterior pole, extending from the egg periphery to the inner yolk mass. At gastrulation the cephalic furrow is shifted toward the anterior and the germband extends all the way to the posterior end. Analysis of germline clones indicates that the torso mutant is germline autonomous (Schupbach and Wieschaus, 1986, Dev. Biol. 113: 443-48).
Recent Updates
Description	What does this section display? This section contains items that were added to this record for each release. It currently only tracks new links between this FlyBase report and other FlyBase data classes (e.g. genes, references, stocks) or controlled vocabulary terms (e.g. GO, anatomy terms). What does this section not display? This section does not currently display links that were removed or gene model changes.
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FB2012_01	Sequence features BKN50419 BKN50746 HFA07707 BKN24507
FB2011_10
All updates	Click here to see a list of all updates to this record from FB2010_08 and on.
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 43E11-43E12
Experimentally Determined Cytological Location
Cytogenetic map Notes References 43D3-43E7   (determined by in situ hybridisation)   (Sprenger et al., 1989) 43E5-43E8   (determined by in situ hybridisation)   (Casanova and Struhl, 1989)
Experimentally Determined Recombination Data
Location	2-57 (Schupbach and Wieschaus, 1989, Tearle and Nusslein-Volhard, 1987)
Left of (cM)	sca (Szabad et al., 1989) cn (Tearle and Nusslein-Volhard, 1987)
Right of (cM)	lt (Szabad et al., 1989)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\tor 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) tor-RA FBtr0088938  NM_057414   3148   923 tor-RB FBtr0112837  NM_001103741   3040   914 tor-RC FBtr0112838  NM_001103742   3212   923
Additional Transcript Data & Comments
Reported size (kB)	5.5, 3.6 (northern blot) (Sprenger et al., 1989)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein tor-PA   FBpp0088012   105.2   923   6.74     NP_476762   AAF59203 tor-PB   FBpp0111750   104.0   914   7.22     NP_001097211   ABV53713 tor-PC   FBpp0111751   105.2   923   6.74     NP_001097212   ABV53714
Additional Polypeptide Data & Comments
Reported size (kDa)	923 (aa); 105 (kD predicted) (Sprenger et al., 1989)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Protein kinase, catalytic domain (IPR000719) Tyrosine-protein kinase, active site (IPR008266) Protein kinase-like domain (IPR011009) Fibronectin, type III (IPR003961) Protein kinase, ATP binding site (IPR017441) Tyrosine-protein kinase, receptor torso (IPR020724) Tyrosine-protein kinase, catalytic domain (IPR020635) Serine-threonine/tyrosine-protein kinase (IPR001245)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA201646   AA246498   AA263625   AA263822   AA264180   AA438904   AA439803   AA540175   AA540330   AA540686   AA541039   AA821010   AA950882   AA951734   AE013599 AAF59203   AE013599 ABV53713   AE013599 ABV53714   AC004283   AI512430   AI513415   AI513890   AI515103   AY071403 AAL49025   BI212478   BI242319   BI243532   BI243609   BI363089   BI364029   BI364057   BI366917   BI367074   BI375973   BT030191   BT125648   BT125750   CO267396   CO293158   CO298209   CO298372   CO325420   CO340172   CZ478045   EC053035   EC053051   EC055787   EC084617   EC086394   EC090395   EC212547   EC245582   EC255810   EC258559   EL886358   EL886467   EL886528   EL886581   EL886635   EV575651   EV577173   EV577192   EV584147   EV584577   GH941402   GH941916   GH942936   GH943233   GH943343   GH943989   GH944502   GH945726   GH946372   GH946593   GH946771   GH947303   GH947710   GH947741   GH949197   GH949280   GH949679   GH950433   GH950920   GH951084   GH951468   GH953562   GH954017   GH954287   GH957322   GH957818   GH959763   GH961363   GH961606   GH962056   GH962234   GH962450   GH962653   GH962711   GH962840   GH964744   GH965471   GH965504   GH966198   GH969996   GH970029   GH970609   GH972747   GH972827   GH973096   GH974074   GH976808   GH980049   GH981809   GH982482   GH982843   GH983395   GH984517   GH984738   X15150 CAA33247
UniProtKB/Swiss-Prot	P18475
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   organism | medial   (Casanova and Struhl, 1993) northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage | early     (Sprenger et al., 1989) adult stage | female     (Sprenger et al., 1989) radioisotope in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage | early   organism | ubiquitous   (Sprenger et al., 1989) adult stage & oogenesis   nurse cell   (Sprenger et al., 1989) germarium   (Sprenger et al., 1989) adult stageoogenesis stage S11 -- S14   oocyte   (Sprenger et al., 1989)
Additional Descriptive Data
	The Kr promoter directs expression of tor mRNA to the central portion of the embryo. (Casanova and Struhl, 1993) tor transcripts are detected mainly in early embryos and in adult females on northern blots. They are detected throughout development at a much lower level. The 5.5kb transcript is present at a significantly lower level than the 3.6kb transcript. (Sprenger et al., 1989) tor transcripts are detected at the earliest stages of oogenesis. They accumulate in the nurse cells until stage S10 after which they are transported to the oocyte. They become evenly distributed between the nurse cells and the oocyte. In early embryos, they are uniformly distributed. During syncytial blastoderm, the RNA moves toward the periphery of the embryos and is mainly situated in the cytoplasm underneath the nuclei. No signal is detected after cellularization. (Sprenger et al., 1989)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
No Assay Recorded Stage Tissue/Position (including subcellular localization) Reference embryonic stage   organism | medial   (Casanova and Struhl, 1993) organism | posterior   (Casanova and Struhl, 1993) immunolocalization Stage Tissue/Position (including subcellular localization) Reference embryonic stage 4     (Casanova and Struhl, 1989)
Additional Descriptive Data
	tor protein is first detected in embryos around the 9th nuclear division at which stage the nuclei have just arrived at the periphery. Protein levels increase over the next several nuclear division cycles prior to the cellularization of the blastoderm. tor protein then decreases over the next couple of hours after which it is no longer detected. (Casanova and Struhl, 1989) The tor protein is expressed in the central portion of the embryo. (Casanova and Struhl, 1993) NRE elements causes tor protein to decline in a graded fashion in the posterior half of the embryo. (Casanova and Struhl, 1993)
Marker for
Subcellular Localization
CV Term	plasma membrane (Casanova and Struhl, 1989)
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 FBgn0003733    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0003733 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 extremely low expression. Peak expression observed within 00-06 hour embryonic stages, in adult female 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 FBgn0003733 expression level Developmental Stage   Expression Level embryo 00-02hr    3891 embryo 02-04hr    2845 embryo 04-06hr    766 embryo 06-08hr    558 embryo 08-10hr    888 embryo 10-12hr    510 embryo 12-14hr    719 embryo 14-16hr    767 embryo 16-18hr    674 embryo 18-20hr    251 embryo 20-22hr    968 embryo 22-24hr    947 larva L1    8 larva L2    18 larva L3 12hr old    13 larva L3 puffstage 1-2    28 larva L3 puffstage 3-6    43 larva L3 puffstage 7-9    56 white prepupae new    55 white prepupae 12hr    98 white prepupae 24hr    79 pupae 2d postWPP    86 pupae 3d postWPP    69 pupae 4d postWPP    36 adult male 01day    70 adult male 05day    109 adult male 30day    80 adult female 01day    979 adult female 05day    2465 adult female 30day    2414 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (3891) embryo 02-04hr  (2845) embryo 04-06hr  (766) embryo 06-08hr  (558) embryo 08-10hr  (888) embryo 10-12hr  (510) embryo 12-14hr  (719) embryo 14-16hr  (767) embryo 16-18hr  (674) embryo 18-20hr  (251) embryo 20-22hr  (968) embryo 22-24hr  (947) larva L1    8 larva L2    18 larva L3 12hr old    13 larva L3 puffstage 1-2    28 larva L3 puffstage 3-6    43 larva L3 puffstage 7-9    56 white prepupae new    55 white prepupae 12hr    98 white prepupae 24hr    79 pupae 2d postWPP    86 pupae 3d postWPP    69 pupae 4d postWPP    36 adult male 01day    70 adult male 05day  (109) adult male 30day    80 adult female 01day  (979) adult female 05day  (2465) adult female 30day  (2414) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (3891) embryo 02-04hr  (2845) embryo 04-06hr    766 embryo 06-08hr    558 embryo 08-10hr    888 embryo 10-12hr    510 embryo 12-14hr    719 embryo 14-16hr    767 embryo 16-18hr    674 embryo 18-20hr    251 embryo 20-22hr    968 embryo 22-24hr    947 larva L1    8 larva L2    18 larva L3 12hr old    13 larva L3 puffstage 1-2    28 larva L3 puffstage 3-6    43 larva L3 puffstage 7-9    56 white prepupae new    55 white prepupae 12hr    98 white prepupae 24hr    79 pupae 2d postWPP    86 pupae 3d postWPP    69 pupae 4d postWPP    36 adult male 01day    70 adult male 05day    109 adult male 30day    80 adult female 01day    979 adult female 05day  (2465) adult female 30day  (2414) Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    3891 embryo 02-04hr    2845 embryo 04-06hr    766 embryo 06-08hr    558 embryo 08-10hr    888 embryo 10-12hr    510 embryo 12-14hr    719 embryo 14-16hr    767 embryo 16-18hr    674 embryo 18-20hr    251 embryo 20-22hr    968 embryo 22-24hr    947 larva L1    8 larva L2    18 larva L3 12hr old    13 larva L3 puffstage 1-2    28 larva L3 puffstage 3-6    43 larva L3 puffstage 7-9    56 white prepupae new    55 white prepupae 12hr    98 white prepupae 24hr    79 pupae 2d postWPP    86 pupae 3d postWPP    69 pupae 4d postWPP    36 adult male 01day    70 adult male 05day    109 adult male 30day    80 adult female 01day    979 adult female 05day    2465 adult female 30day    2414 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    3891 embryo 02-04hr    2845 embryo 04-06hr    766 embryo 06-08hr    558 embryo 08-10hr    888 embryo 10-12hr    510 embryo 12-14hr    719 embryo 14-16hr    767 embryo 16-18hr    674 embryo 18-20hr    251 embryo 20-22hr    968 embryo 22-24hr    947 larva L1    8 larva L2    18 larva L3 12hr old    13 larva L3 puffstage 1-2    28 larva L3 puffstage 3-6    43 larva L3 puffstage 7-9    56 white prepupae new    55 white prepupae 12hr    98 white prepupae 24hr    79 pupae 2d postWPP    86 pupae 3d postWPP    69 pupae 4d postWPP    36 adult male 01day    70 adult male 05day    109 adult male 30day    80 adult female 01day    979 adult female 05day    2465 adult female 30day    2414 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0003733 expression level Developmental Stage   Expression Level embryo 00-02hr    3891 embryo 02-04hr    2845 embryo 04-06hr    766 embryo 06-08hr    558 embryo 08-10hr    888 embryo 10-12hr    510 embryo 12-14hr    719 embryo 14-16hr    767 embryo 16-18hr    674 embryo 18-20hr    251 embryo 20-22hr    968 embryo 22-24hr    947 larva L1    8 larva L2    18 larva L3 12hr old    13 larva L3 puffstage 1-2    28 larva L3 puffstage 3-6    43 larva L3 puffstage 7-9    56 white prepupae new    55 white prepupae 12hr    98 white prepupae 24hr    79 pupae 2d postWPP    86 pupae 3d postWPP    69 pupae 4d postWPP    36 adult male 01day    70 adult male 05day    109 adult male 30day    80 adult female 01day    979 adult female 05day    2465 adult female 30day    2414 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (3891) embryo 02-04hr  (2845) embryo 04-06hr  (766) embryo 06-08hr  (558) embryo 08-10hr  (888) embryo 10-12hr  (510) embryo 12-14hr  (719) embryo 14-16hr  (767) embryo 16-18hr  (674) embryo 18-20hr  (251) embryo 20-22hr  (968) embryo 22-24hr  (947) larva L1    8 larva L2    18 larva L3 12hr old    13 larva L3 puffstage 1-2    28 larva L3 puffstage 3-6    43 larva L3 puffstage 7-9    56 white prepupae new    55 white prepupae 12hr    98 white prepupae 24hr    79 pupae 2d postWPP    86 pupae 3d postWPP    69 pupae 4d postWPP    36 adult male 01day    70 adult male 05day  109 adult male 30day    80 adult female 01day  (979) adult female 05day  (2465) adult female 30day  (2414) Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (3891) embryo 02-04hr  (2845) embryo 04-06hr    766 embryo 06-08hr    558 embryo 08-10hr    888 embryo 10-12hr    510 embryo 12-14hr    719 embryo 14-16hr    767 embryo 16-18hr    674 embryo 18-20hr    251 embryo 20-22hr    968 embryo 22-24hr    947 larva L1    8 larva L2    18 larva L3 12hr old    13 larva L3 puffstage 1-2    28 larva L3 puffstage 3-6    43 larva L3 puffstage 7-9    56 white prepupae new    55 white prepupae 12hr    98 white prepupae 24hr    79 pupae 2d postWPP    86 pupae 3d postWPP    69 pupae 4d postWPP    36 adult male 01day    70 adult male 05day    109 adult male 30day    80 adult female 01day    979 adult female 05day  2465 adult female 30day  2414 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    3891 embryo 02-04hr    2845 embryo 04-06hr    766 embryo 06-08hr    558 embryo 08-10hr    888 embryo 10-12hr    510 embryo 12-14hr    719 embryo 14-16hr    767 embryo 16-18hr    674 embryo 18-20hr    251 embryo 20-22hr    968 embryo 22-24hr    947 larva L1    8 larva L2    18 larva L3 12hr old    13 larva L3 puffstage 1-2    28 larva L3 puffstage 3-6    43 larva L3 puffstage 7-9    56 white prepupae new    55 white prepupae 12hr    98 white prepupae 24hr    79 pupae 2d postWPP    86 pupae 3d postWPP    69 pupae 4d postWPP    36 adult male 01day    70 adult male 05day    109 adult male 30day    80 adult female 01day    979 adult female 05day    2465 adult female 30day    2414 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    3891 embryo 02-04hr    2845 embryo 04-06hr    766 embryo 06-08hr    558 embryo 08-10hr    888 embryo 10-12hr    510 embryo 12-14hr    719 embryo 14-16hr    767 embryo 16-18hr    674 embryo 18-20hr    251 embryo 20-22hr    968 embryo 22-24hr    947 larva L1    8 larva L2    18 larva L3 12hr old    13 larva L3 puffstage 1-2    28 larva L3 puffstage 3-6    43 larva L3 puffstage 7-9    56 white prepupae new    55 white prepupae 12hr    98 white prepupae 24hr    79 pupae 2d postWPP    86 pupae 3d postWPP    69 pupae 4d postWPP    36 adult male 01day    70 adult male 05day    109 adult male 30day    80 adult female 01day    979 adult female 05day    2465 adult female 30day    2414 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 FBgn0003733    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0003733 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: adult ovary. [download data (TSV)] Guide to FlyAtlas expression level colors   No expression (0 - 9.999)   Low expression (10 - 99.999)   Moderate expression (100 - 499.999)   High level expression (500 - 999.999)   Very high expression (1000 - 25000) Linear, scaled to maximum FBgn0003733 expression level Tissue   Expression Level Larval Central Nervous System    2.9 Larval Midgut    2.4 Larval Hindgut    0.6 Larval Malpighian Tubules    0.9 Larval Fat Body    1.7 Larval Salivary Gland    4.7 Larval Trachea    0.55 Larval Carcass    2.45 Adult Head    0.8 Adult Eye    2.725 Adult Brain    0.8 Adult Thoracic-Abdominal Ganglion    1 Adult Crop    2.1 Adult Midgut    1.6 Adult Hindgut    2.2 Adult Malpighian Tubules    2.8 Adult Fat Body    61.4 Adult Salivary Gland    3.9 Adult Heart    45.525 Adult VirginFemale Spermatheca    6.1 Adult InseminatedFemale Spermatheca    2.8 Adult Ovary    291.9 Adult Testis    4.1 Adult Male Accessory Gland    2.2 Adult Carcass    63.5 Expression Level Scale  None   Low   Moderate  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    2.9 Larval Midgut    2.4 Larval Hindgut    0.6 Larval Malpighian Tubules    0.9 Larval Fat Body    1.7 Larval Salivary Gland    4.7 Larval Trachea    0.55 Larval Carcass    2.45 Adult Head    0.8 Adult Eye    2.725 Adult Brain    0.8 Adult Thoracic-Abdominal Ganglion    1 Adult Crop    2.1 Adult Midgut    1.6 Adult Hindgut    2.2 Adult Malpighian Tubules    2.8 Adult Fat Body    61.4 Adult Salivary Gland    3.9 Adult Heart    45.525 Adult VirginFemale Spermatheca    6.1 Adult InseminatedFemale Spermatheca    2.8 Adult Ovary    291.9 Adult Testis    4.1 Adult Male Accessory Gland    2.2 Adult Carcass    63.5 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    2.9 Larval Midgut    2.4 Larval Hindgut    0.6 Larval Malpighian Tubules    0.9 Larval Fat Body    1.7 Larval Salivary Gland    4.7 Larval Trachea    0.55 Larval Carcass    2.45 Adult Head    0.8 Adult Eye    2.725 Adult Brain    0.8 Adult Thoracic-Abdominal Ganglion    1 Adult Crop    2.1 Adult Midgut    1.6 Adult Hindgut    2.2 Adult Malpighian Tubules    2.8 Adult Fat Body    61.4 Adult Salivary Gland    3.9 Adult Heart    45.525 Adult VirginFemale Spermatheca    6.1 Adult InseminatedFemale Spermatheca    2.8 Adult Ovary    291.9 Adult Testis    4.1 Adult Male Accessory Gland    2.2 Adult Carcass    63.5 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    2.9 Larval Midgut    2.4 Larval Hindgut    0.6 Larval Malpighian Tubules    0.9 Larval Fat Body    1.7 Larval Salivary Gland    4.7 Larval Trachea    0.55 Larval Carcass    2.45 Adult Head    0.8 Adult Eye    2.725 Adult Brain    0.8 Adult Thoracic-Abdominal Ganglion    1 Adult Crop    2.1 Adult Midgut    1.6 Adult Hindgut    2.2 Adult Malpighian Tubules    2.8 Adult Fat Body    61.4 Adult Salivary Gland    3.9 Adult Heart    45.525 Adult VirginFemale Spermatheca    6.1 Adult InseminatedFemale Spermatheca    2.8 Adult Ovary    291.9 Adult Testis    4.1 Adult Male Accessory Gland    2.2 Adult Carcass    63.5 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to maximum FBgn0003733 expression level Tissue   Expression Level Larval Central Nervous System    2.9 Larval Midgut    2.4 Larval Hindgut    0.6 Larval Malpighian Tubules    0.9 Larval Fat Body    1.7 Larval Salivary Gland    4.7 Larval Trachea    0.55 Larval Carcass    2.45 Adult Head    0.8 Adult Eye    2.725 Adult Brain    0.8 Adult Thoracic-Abdominal Ganglion    1 Adult Crop    2.1 Adult Midgut    1.6 Adult Hindgut    2.2 Adult Malpighian Tubules    2.8 Adult Fat Body    61.4 Adult Salivary Gland    3.9 Adult Heart    45.525 Adult VirginFemale Spermatheca    6.1 Adult InseminatedFemale Spermatheca    2.8 Adult Ovary    291.9 Adult Testis    4.1 Adult Male Accessory Gland    2.2 Adult Carcass    63.5 Expression Level Scale  None   Low   Moderate   High  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    2.9 Larval Midgut    2.4 Larval Hindgut    0.6 Larval Malpighian Tubules    0.9 Larval Fat Body    1.7 Larval Salivary Gland    4.7 Larval Trachea    0.55 Larval Carcass    2.45 Adult Head    0.8 Adult Eye    2.725 Adult Brain    0.8 Adult Thoracic-Abdominal Ganglion    1 Adult Crop    2.1 Adult Midgut    1.6 Adult Hindgut    2.2 Adult Malpighian Tubules    2.8 Adult Fat Body    61.4 Adult Salivary Gland    3.9 Adult Heart    45.525 Adult VirginFemale Spermatheca    6.1 Adult InseminatedFemale Spermatheca    2.8 Adult Ovary    291.9 Adult Testis    4.1 Adult Male Accessory Gland    2.2 Adult Carcass    63.5 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    2.9 Larval Midgut    2.4 Larval Hindgut    0.6 Larval Malpighian Tubules    0.9 Larval Fat Body    1.7 Larval Salivary Gland    4.7 Larval Trachea    0.55 Larval Carcass    2.45 Adult Head    0.8 Adult Eye    2.725 Adult Brain    0.8 Adult Thoracic-Abdominal Ganglion    1 Adult Crop    2.1 Adult Midgut    1.6 Adult Hindgut    2.2 Adult Malpighian Tubules    2.8 Adult Fat Body    61.4 Adult Salivary Gland    3.9 Adult Heart    45.525 Adult VirginFemale Spermatheca    6.1 Adult InseminatedFemale Spermatheca    2.8 Adult Ovary    291.9 Adult Testis    4.1 Adult Male Accessory Gland    2.2 Adult Carcass    63.5 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    2.9 Larval Midgut    2.4 Larval Hindgut    0.6 Larval Malpighian Tubules    0.9 Larval Fat Body    1.7 Larval Salivary Gland    4.7 Larval Trachea    0.55 Larval Carcass    2.45 Adult Head    0.8 Adult Eye    2.725 Adult Brain    0.8 Adult Thoracic-Abdominal Ganglion    1 Adult Crop    2.1 Adult Midgut    1.6 Adult Hindgut    2.2 Adult Malpighian Tubules    2.8 Adult Fat Body    61.4 Adult Salivary Gland    3.9 Adult Heart    45.525 Adult VirginFemale Spermatheca    6.1 Adult InseminatedFemale Spermatheca    2.8 Adult Ovary    291.9 Adult Testis    4.1 Adult Male Accessory Gland    2.2 Adult Carcass    63.5 Expression Level Scale  None   Low   Moderate   High   Very high  Heatmap Tissue   Expression Level Larval Central Nervous System     Larval Midgut     Larval Hindgut     Larval Malpighian Tubules     Larval Fat Body     Larval Salivary Gland     Larval Trachea     Larval Carcass     Adult Head     Adult Eye     Adult Brain     Adult Thoracic-Abdominal Ganglion     Adult Crop     Adult Midgut     Adult Hindgut     Adult Malpighian Tubules     Adult Fat Body     Adult Salivary Gland     Adult Heart     Adult VirginFemale Spermatheca     Adult InseminatedFemale Spermatheca     Adult Ovary     Adult Testis     Adult Male Accessory Gland     Adult Carcass     FlyAtlas Organ/Tissue Expression, larval vs. adult Larval Expression Level Tissue Adult Expression Level   NA  Head    0.8   NA  Eye    2.725   NA  Brain    0.8   2.9  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    1   NA  Crop    2.1   2.4  Midgut    1.6   0.6  Hindgut    2.2   0.9  Malpighian Tubules    2.8   1.7  Fat Body    61.4   4.7  Salivary Gland    3.9   NA  Heart    45.525   0.55  Trachea    NA   NA  VirginFemale Spermatheca    6.1   NA  InseminatedFemale Spermatheca    2.8   NA  Ovary    291.9   NA  Testis    4.1   NA  Male Accessory Gland    2.2   2.45  Carcass    63.5 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_01
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003733 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG1389-RA FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for tor Stages(s) 1-3 Stages(s) 4-6 Stages(s) 7-8 Stages(s) 9-10
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal torQuad torQuint torY767+772F lethal | cold sensitive tor12D tor13D lethal | dominant | maternal effect | heat sensitive tor11D lethal | embryonic stage | germline clone | non-rescuable maternal effect | recessive tor2R-51-6 tor2R-70-6 tor2R-145-24 tor2R-219-37 tor2R-255-40 lethal | embryonic stage | maternal effect | cold sensitive tor12D lethal | embryonic stage | maternal effect | recessive tor2 tor3 tor4 tor5 tor6 tor7 tor8 tor9 tor11D lethal | embryonic stage | maternal effect | recessive | heat sensitive tor11D lethal | gap | heat sensitive tor11D lethal | heat sensitive, with Scer\GAL4Bx-MS1096 tor4021.Scer\UAS lethal | maternal effect tor11D lethal | maternal effect | recessive tor10 tor14 torrv1 torrv2 torrv3 torrv4 torrv5 torrv6 torrv7 torrv8 torrv9 torrv10 torrv11 torrv12 torrv13 torrv14 torrv15 torrv16 torrv17 torrv18 torrv19 torrv20 torrv21 torrv22 torrv23 torrv24 torrv25 torrv26 torrv27 torrv28 torrv29 torrv30 torrv31 torrv32 torrv33 torrv34 torrv35 torrv36 torrv37 torrv38 torrv39 torrv40 torrv41 torrv42 torrv43 torrv44 torrv45 torrv46 torrv47 torrv48 torrv49 torrv50 torrv51 torrv52 torrv53 torrv54 torrv55 torrv56 torrv57 torrv58 torrv59 torrv60 torrv61 torrv62 torrv63 torrv64 torrv65 torrv66 lethal | recessive tor4 semi-lethal torY630F viable tore00150 viable, with Scer\GAL4pnr-MD237 torGD117 torGD2613 torGD14396 Sterility Allele female sterile | dominant tor13D female sterile | heat sensitive tor11D female sterile | recessive tor2 tor3 tor4 tor5 tor6 tor7 tor8 tor9 tor11D female sterile germline-dependent | dominant tor13D fertile tore00150 Other Phenotypes Allele developmental rate defective, with Scer\GAL4phm.PO torGD14396 developmental rate defective | dominant | heat sensitive tor11D developmental rate defective | recessive torunspecified hyperactive | dominant tor13D hypoactive | dominant tor13D large body, with Scer\GAL4phm.PO torGD14396 large body | recessive torunspecified small body | dominant | male limited | heat sensitive tor11D visible, with Scer\GAL4Bx-MS1096 tor4021.Scer\UAS visible | heat sensitive tor11D wild-type tor+t11.5 torBtor torhb1+hb2.96-1 torY644F torY656F torY698F Phenotype manifest in Allele abdomen | posterior torrv66 abdominal 8 ventral denticle belt | maternal effect torrv66 abdominal 11 anal tuft | embryonic stage torrv66 abdominal segment tor11D abdominal segment 5 tor1 tor2 abdominal segment 6 tor1 tor2 abdominal segment 7 tor1 tor2 abdominal segment 8 tor1 tor2 tor4 abdominal segment 9 tor1 tor2 tor4 abdominal segment 10 tor1 tor2 tor4 abdominal segment 11 tor4 adult labral sense organ tor4 anterior midgut primordium | maternal effect | heat sensitive tor11D cephalic furrow tor1 tor2 cephalic furrow | maternal effect tor2 tor3 tor4 tor5 tor6 tor7 tor8 tor9 cephalic furrow | maternal effect | heat sensitive tor11D cephalopharyngeal skeleton tor1 cephalopharyngeal skeleton | maternal effect | cold sensitive tor12D denticle belt tor12D denticle belt | maternal effect tor12D denticle belt | maternal effect | cold sensitive tor12D denticle belt | maternal effect | heat sensitive tor11D dorsal bridge torunspecified dorsopharyngeal organ tor4 embryo | germline clone | maternal effect | terminal tor2R-145-24 tor2R-219-37 tor2R-255-40 tor2R-51-6 tor2R-70-6 embryo | posterior tor3 torRX embryo | terminal tor11D embryonic/first instar larval cuticle tor13D torrv66 embryonic/first instar larval cuticle | maternal effect tor12D tor13D tor2 tor3 tor4 tor5 tor6 tor7 tor8 tor9 embryonic/first instar larval cuticle | maternal effect (with tor12D) torrv66 embryonic/first instar larval cuticle | maternal effect (with torrv66) tor12D embryonic/first instar larval cuticle | maternal effect | cold sensitive tor12D embryonic/first instar larval cuticle | maternal effect | heat sensitive tor11D embryonic/larval anus | embryonic stage tor1 embryonic/larval brain | embryonic stage tor4 embryonic/larval esophagus | embryonic stage tor4 embryonic/larval labral nerve tor4 embryonic/larval labral segment sense organ tor4 embryonic/larval midgut | embryonic stage | posterior torrv66 embryonic/larval optic lobe | embryonic stage tor4 embryonic/larval posterior spiracle | embryonic stage tor1 embryonic abdominal segment 2 | heat sensitive tor11D embryonic abdominal segment 3 | heat sensitive tor11D embryonic abdominal segment 4 | heat sensitive tor11D embryonic abdominal segment 4 | maternal effect tor11D embryonic abdominal segment 5 | heat sensitive tor11D embryonic abdominal segment 7 torunspecified embryonic abdominal segment 8 tor3 torrv66 torunspecified embryonic abdominal segment | gap | heat sensitive tor11D embryonic abdominal segment | maternal effect tor12D embryonic abdominal segment | maternal effect | heat sensitive tor11D embryonic anal pad torrv66 embryonic epidermis torQuad torQuint torTCE.Asp718 torTCE.SpeI torY630F torY767+772F torY918F torhb1+hb2.1-96 torhb1+hb2.96-1 embryonic head tor4 torrv66 embryonic head | heat sensitive tor11D embryonic head | maternal effect | posterior | heat sensitive tor11D embryonic labial segment torunspecified embryonic labral segment | maternal effect tor2 tor3 tor4 tor5 tor6 tor7 tor8 tor9 embryonic telson torrv66 embryonic thoracic segment | gap | heat sensitive tor11D embryonic thoracic segment | maternal effect | heat sensitive tor11D endoderm tor13D extended germ band embryo | maternal effect tor2 tor3 tor4 tor5 tor6 tor7 tor8 tor9 filzkorper tor1 tor12D torrv66 filzkorper | ectopic | maternal effect tor12D filzkorper | embryonic stage torrv66 filzkorper | maternal effect | cold sensitive tor12D gastrula embryo torrv66 germline cell tor+t11.5 head tor4 labral segment tor4 labrum tor1 tor2 labrum | embryonic stage torrv66 ocular segment tor1 tor2 pharyngeal monoscolopidial chordotonal organ tor4 pole cell | ectopic | maternal effect torrv66 posterior midgut primordium torrv66 posterior midgut primordium | maternal effect tor2 tor3 tor4 tor5 tor6 tor7 tor8 tor9 torrv66 pupa, with Scer\GAL4phm.PO torGD14396 salivary gland common duct primordium | maternal effect torrv66 salivary gland common duct primordium | maternal effect | heat sensitive tor11D stomatogastric nervous system tor4 stomatogastric nervous system | embryonic stage tor4 telson tor1 tor2 tor4 torunspecified telson | maternal effect tor2 tor3 tor4 tor5 tor6 tor7 tor8 tor9 ventral furrow tor1 tor2 ventral furrow | maternal effect | heat sensitive tor11D visceral mesoderm tor13D wing, with Scer\GAL4Bx-MS1096 tor4021.Scer\UAS
Classical Alleles ( 93 )
For All Classical Alleles Show
Allele of tor Class Mutagen Stocks Known lesion tor1 loss of function allele ethyl methanesulfonate 7 Yes tor4 loss of function allele, amorphic allele - genetic evidence, gain of function allele ethyl methanesulfonate 2 Yes tor11D hypermorphic allele - genetic evidence, antimorphic allele - genetic evidence, loss of function allele, gain of function allele ethyl methanesulfonate 2 Yes tore00150 piggyBac transposase 2 -- torNP3370 2 -- tor8 amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes torrv66 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate X ray 0 Yes tor10 ethyl methanesulfonate 0 -- tor12D antimorphic allele - genetic evidence, gain of function allele ethyl methanesulfonate 0 Yes tor13D antimorphic allele - genetic evidence, loss of function allele, gain of function allele ethyl methanesulfonate 0 Yes tor14 loss of function allele PM hybrid dysgenesis 0 Yes tor2 loss of function allele ethyl methanesulfonate 0 Yes tor2R-145-24 ethyl methanesulfonate 0 -- tor2R-219-37 ethyl methanesulfonate 0 -- tor2R-255-40 ethyl methanesulfonate 0 -- tor2R-51-6 ethyl methanesulfonate 0 -- tor2R-70-6 ethyl methanesulfonate 0 -- tor3 ethyl methanesulfonate 0 Yes tor5 ethyl methanesulfonate 0 Yes tor6 ethyl methanesulfonate 0 Yes tor7 ethyl methanesulfonate 0 -- tor9 loss of function allele ethyl methanesulfonate 0 Yes torgf gain of function allele 0 -- torQK45 loss of function allele ethyl methanesulfonate 0 Yes torrv10 ethyl methanesulfonate 0 -- torrv11 ethyl methanesulfonate 0 -- torrv12 ethyl methanesulfonate 0 -- torrv13 ethyl methanesulfonate 0 -- torrv14 ethyl methanesulfonate 0 -- torrv15 ethyl methanesulfonate 0 -- torrv16 ethyl methanesulfonate 0 -- torrv17 ethyl methanesulfonate 0 -- torrv18 ethyl methanesulfonate 0 -- torrv19 ethyl methanesulfonate 0 -- torrv1 ethyl methanesulfonate 0 -- torrv20 ethyl methanesulfonate 0 -- torrv21 ethyl methanesulfonate 0 -- torrv22 ethyl methanesulfonate 0 -- torrv23 ethyl methanesulfonate 0 -- torrv24 ethyl methanesulfonate 0 -- torrv25 ethyl methanesulfonate 0 -- torrv26 ethyl methanesulfonate 0 -- torrv27 ethyl methanesulfonate 0 -- torrv28 ethyl methanesulfonate 0 -- torrv29 ethyl methanesulfonate 0 -- torrv2 ethyl methanesulfonate 0 -- torrv30 ethyl methanesulfonate 0 -- torrv31 ethyl methanesulfonate 0 -- torrv32 ethyl methanesulfonate 0 -- torrv33 ethyl methanesulfonate 0 -- torrv34 ethyl methanesulfonate 0 -- torrv35 ethyl methanesulfonate 0 -- torrv36 ethyl methanesulfonate 0 -- torrv37 ethyl methanesulfonate 0 -- torrv38 ethyl methanesulfonate 0 -- torrv39 ethyl methanesulfonate 0 -- torrv3 ethyl methanesulfonate 0 -- torrv40 ethyl methanesulfonate 0 -- torrv41 ethyl methanesulfonate 0 -- torrv42 ethyl methanesulfonate 0 -- torrv43 ethyl methanesulfonate 0 -- torrv44 ethyl methanesulfonate 0 -- torrv45 ethyl methanesulfonate 0 -- torrv46 ethyl methanesulfonate 0 -- torrv47 ethyl methanesulfonate 0 -- torrv48 ethyl methanesulfonate 0 -- torrv49 ethyl methanesulfonate 0 -- torrv4 ethyl methanesulfonate 0 -- torrv50 ethyl methanesulfonate 0 -- torrv51 ethyl methanesulfonate 0 -- torrv52 ethyl methanesulfonate 0 -- torrv53 ethyl methanesulfonate 0 -- torrv54 ethyl methanesulfonate 0 -- torrv55 ethyl methanesulfonate 0 -- torrv56 ethyl methanesulfonate 0 -- torrv57 ethyl methanesulfonate 0 -- torrv58 ethyl methanesulfonate 0 -- torrv59 ethyl methanesulfonate 0 -- torrv5 ethyl methanesulfonate 0 -- torrv60 ethyl methanesulfonate 0 -- torrv61 ethyl methanesulfonate 0 -- torrv62 X ray ethyl methanesulfonate 0 -- torrv63 X ray ethyl methanesulfonate 0 -- torrv64 X ray ethyl methanesulfonate 0 -- torrv65 ethyl methanesulfonate 0 -- torrv67 ethyl methanesulfonate X ray 0 Yes torrv6 ethyl methanesulfonate 0 -- torrv7 ethyl methanesulfonate 0 -- torrv8 ethyl methanesulfonate 0 -- torrv9 ethyl methanesulfonate 0 -- torRX 0 -- torTC 0 -- torunspecified 0 --
Alleles Carried on Transgenic Constructs ( 44 )
For All Alleles Carried on Transgenic Constructs Show
Allele of tor Class Mutagen Stocks Known lesion torGD2613 in vitro construct - RNAi in vitro construct - regulatory fusion 3 Yes torGD14396 in vitro construct - RNAi in vitro construct - regulatory fusion 2 Yes torGD117 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes torGL00222 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes torGLV21002 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes torHMS00021 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes torKK105371 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes tor+t11.5 in vitro construct - genomic fragment 0 Yes tor13D.hs.sev in vitro construct - regulatory fusion 0 Yes tor4021.Scer\UAS in vitro construct - regulatory fusion 0 Yes tora.cFa in vitro construct 0 Yes toract.RL3 in vitro construct 0 Yes torBHM in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes torBTKM in vitro construct - amino acid replacement 0 Yes torBTKMCy in vitro construct - deletion 0 Yes torBTKMd69 in vitro construct - deletion 0 Yes torBtor4021 in vitro construct - amino acid replacement 0 Yes torBtor4021TKM in vitro construct - amino acid replacement 0 Yes torBtor in vitro construct - regulatory fusion 0 Yes torBtord69 in vitro construct - deletion 0 Yes torBtordB in vitro construct - deletion 0 Yes torBtorY9 in vitro construct - amino acid replacement 0 Yes torBtorY9TKM in vitro construct - amino acid replacement 0 Yes torGOF.cLa in vitro construct 0 Yes torhb1+hb2.1-96 in vitro construct - regulatory fusion 0 Yes torhb1+hb2.96-1 in vitro construct - regulatory fusion 0 Yes torhb1+hb2 in vitro construct - regulatory fusion 0 Yes torhb2+hb2 in vitro construct - regulatory fusion 0 Yes torKr.P in vitro construct - regulatory fusion 0 Yes torNRE hypomorphic allele - genetic evidence in vitro construct - regulatory fusion 0 Yes torQuad in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes torQuint in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes torScer\UAS.P\T.cGa in vitro construct - regulatory fusion 0 Yes torTCE.Asp718 in vitro construct - regulatory fusion 0 Yes torTCE.SpeI in vitro construct - regulatory fusion 0 Yes torTM.hs.T:Scer\GAL4,T:Hsim\VP16,T:Hsap\MYC in vitro construct 0 Yes torWT.cLa in vitro construct 0 Yes torY630F loss of function allele in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes torY644F in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes torY656F in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes torY698F in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes torY767+772F in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes torY918F gain of function allele in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes torYY630:918FF in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2R)cn-S5 (Heitzler et al., 1993) Df(2R)cn-S6 (Heitzler et al., 1993) Df(2R)cn-S8 (Heitzler et al., 1993) Df(2R)Drlrv7 (Heitzler et al., 1993) Df(2R)Drlrv18 (Heitzler et al., 1993) Df(2R)Drlrv24 (Heitzler et al., 1993) Df(2R)Drlrv26 (Heitzler et al., 1993) Df(2R)Exel6054 (Ryder, 2004.4.26) Df(2R)H23 (Tearle and Nusslein-Volhard, 1987) Df(2R)NCX8 (Heitzler et al., 1993) Df(2R)NCX9 (Heitzler et al., 1993) Df(2R)NCX11 (Heitzler et al., 1993) Df(2R)P32a (Schupbach and Wieschaus, 1989, Schupbach and Wieschaus, 1991) Df(2R)pk78s (Schupbach and Wieschaus, 1986, Schupbach and Wieschaus, 1989, Schupbach and Wieschaus, 1991) Df(2R)sple-N3 (Heitzler et al., 1993) Df(2R)ST1 (Schupbach and Wieschaus, 1989, Schupbach and Wieschaus, 1991) Df(2R)tor-rx5 (Tearle and Nusslein-Volhard, 1987)
Not disrupted in	Df(2R)CA53 (Tearle and Nusslein-Volhard, 1987) Df(2R)cos-2 (Heitzler et al., 1993) Df(2R)cos-3 (Heitzler et al., 1993) Df(2R)Drlrv3 (Heitzler et al., 1993) Df(2R)Drlrv5 (Heitzler et al., 1993) Df(2R)Drlrv15 (Heitzler et al., 1993) Df(2R)Drlrv17 (Heitzler et al., 1993) Df(2R)Drlrv20 (Heitzler et al., 1993) Df(2R)Drlrv21 (Heitzler et al., 1993) Df(2R)Drlrv22 (Heitzler et al., 1993) Df(2R)Drlrv25 (Heitzler et al., 1993) Df(2R)Drlrv28 (Heitzler et al., 1993) Df(2R)Drlrv30 (Heitzler et al., 1993) Df(2R)DX1 (Heitzler et al., 1993) Df(2R)DX14 (Heitzler et al., 1993) Df(2R)EW60 (Heitzler et al., 1993) Df(2R)H23 (Heitzler et al., 1993) Df(2R)nap2 (Heitzler et al., 1993) Df(2R)NCX5 (Heitzler et al., 1993) Df(2R)NCX10 (Heitzler et al., 1993) Df(2R)pk-N5 (Heitzler et al., 1993) Df(2R)pwn5 (Heitzler et al., 1993) Df(2R)sple-D1 (Heitzler et al., 1993) Df(2R)sple-D2 (Heitzler et al., 1993) Df(2R)sple-D3 (Heitzler et al., 1993) Df(2R)sple-J1 (Heitzler et al., 1993) Df(2R)sple-J2 (Heitzler et al., 1993)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD117} NA P{GD2613} NA P{GD14396} NA P{KK105371} NA P{TRiP.GL00222} NA P{TRiP.GLV21002} NA P{TRiP.HMS00021} NA P{UASp-tor.G} NA P{UAS-put.tor.N} NA P{UAS-tkv.tor.N} NA P{UAS-tor4021} NA heat-shock construct P{hs-1XMyc-TMTor-GV} NA P{sE-torso4021} NA reporter construct P{tor-LacZ.G} No characterization construct P{hs(FRT.y+)-N-TMTor-GV.3} NA P{hs(FRT.y+)-N-Tor-GV.3} NA P{hs(-FRT)-N-TMTor-GV.3} NA P{hs(-FRT)-N-Tor-GV.3} NA P{Hsp83-pll::tor.4021.bcd.3'UTR} NA P{Hsp83-pll::tor.bcd.3'UTR} NA P{hsp-torso4021-sev} NA P{hs-torso4021-dos-GFP} NA P{sE-KDN.T} NA P{sE-raftor4021} NA P{sE-raftor4021KV} NA P{sE-raftor} NA P{sE-raftorY9} NA P{sE-tor[4021]-ksr.K>M} NA P{sE-tor4021-btl} NA P{sE-tor4021-ksr.fl} NA P{sE-torso4021-DER} NA P{sE-torso4021-DFGF-R2} NA P{sE-torso4021-sev} NA P{sE-torso4021-sevK2242M} NA P{sE-torso4021-sevY2546F} NA P{sE-torso+-sev} NA P{sE-torsoY9-sev} NA P{sev-Raftor9} NA P{tor1-96} NA P{tor96-1} NA P{tor.RL3} NA P{tor(TCE)Asp718} NA P{tor(TCE)SpeI} NA P{tor4021pelle3'} NA P{tor4021pelle} NA P{tor4021tube} NA P{tor+.C} NA P{tor+.S} NA P{torhb1+hb2} NA P{torhb2+hb2} NA P{torKr.P} NA P{torNRE} NA P{torQuad.G} NA P{torQuint.G} NA P{torY630F} NA P{torY644F.G} NA P{torY656F.G} NA P{torY698F.G} NA P{torY767+772F.G} NA P{torY918F} NA P{torYY630/918FF} NA P{tor-Egfr.G} NA P{tor-sev.G} NA P{torso4021-Notch} NA P{UAS-btl::tor.D} NA P{UAS-htl::tor.D} NA P{UAS-mys::tor.D.A834P.T825I.S836N.T837N} NA P{UAS-mys::tor.D.D807V.E810A} NA P{UAS-mys::tor.D.F811A.F814L.E817V} NA P{UAS-mys::tor.D.N828A} NA P{UAS-mys::tor.D.N840A} NA P{UAS-mys::tor.D.P829A} NA P{UAS-mys::tor.D.P841A} NA P{UAS-mys::tor.D.T835A.S836A.T837A} NA P{UAS-mys::tor.D} NA P{UAS-mys::tor.WT} NA P{UAS-pio::tor.B} NA P{UAS-RaftorY9} NA P{UAS-tor4021-EGFR} NA P{UAS-tor4021-htl.MYC} NA P{UAS-tor-HA-CycA} NA P{UAS-torso4021-btl} NA P{UAS-torso4021-htl} NA
Insertions
	Type of insertions Name Expression data miscellaneous insertions P{}tor14 NA PBac{RB}tore00150 NA insertion of enhancer trap binary system P{GawB}torNP3370 No
Gene Ontology: Function, Process & Cellular Component ( 15 unique terms )
Terms Based on Experimental Evidence ( 7 terms )
Molecular Function
CV term Evidence References
protein tyrosine kinase activity inferred from direct assay (Sprenger et al., 1993)
Biological Process
CV term Evidence References
chorion-containing eggshell pattern formation inferred from mutant phenotype (Schupbach and Wieschaus, 1991) gastrulation inferred from mutant phenotype (Li et al., 2003) metamorphosis inferred from mutant phenotype (Rewitz et al., 2009) pole cell migration inferred from mutant phenotype (Li et al., 2003) protein autophosphorylation inferred from direct assay (Sprenger et al., 1993) terminal region determination inferred from mutant phenotype (Sprenger et al., 1989, Jimenez et al., 2000)
Cellular Component ( 0 terms)
Terms Based on Predictions or Assertions ( 10 terms )
Molecular Function
CV term Evidence References
ATP binding inferred from electronic annotation with InterPro:IPR000719, InterPro:IPR017441 (FlyBase Curators et al., 2004-) protein tyrosine kinase activity non-traceable author statement (Morrison et al., 2000) transmembrane receptor protein tyrosine kinase activity inferred from sequence or structural similarity (Sprenger et al., 1989)
Biological Process
CV term Evidence References
anterior/posterior axis specification, embryo traceable author statement (Baek and Lee, 1999) cell fate determination traceable author statement (Bier, 1998) Malpighian tubule morphogenesis traceable author statement (Murakami et al., 1999) protein phosphorylation non-traceable author statement (Morrison et al., 2000, Baek and Lee, 1999) terminal region determination non-traceable author statement (Casci, 2000) torso signaling pathway non-traceable author statement (Swiss-Prot Project Members, 1990.11.1) traceable author statement (Baek and Lee, 1999)
Cellular Component
CV term Evidence References
plasma membrane non-traceable author statement (Murakami et al., 1999)
Sequence Ontology: Class of Gene
	nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
Protein-protein
Interacting group Assay References
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions tor allele Gene References tor3 sty (Casci et al., 1999) tor4 Dsor1 (Lim et al., 1997) tor11D bowl (Doyle and Bishop, 1993) drk (Doyle and Bishop, 1993) Dsor1 (Lim et al., 1999) Ras85D (Doyle and Bishop, 1993) rl (Brunner et al., 1994, Lim et al., 1999) Sos (Doyle and Bishop, 1993) Stat92E (Li et al., 2002) Su(tor)1-1 (Doyle and Bishop, 1993) Su(tor)1-106 (Doyle and Bishop, 1993) Su(tor)1-163 (Doyle and Bishop, 1993) Su(tor)2-3 (Doyle and Bishop, 1993) Su(tor)2-4 (Doyle and Bishop, 1993) Su(tor)2-15 (Doyle and Bishop, 1993) Su(tor)2-48 (Doyle and Bishop, 1993) Su(tor)2-56 (Doyle and Bishop, 1993) Su(tor)2-87 (Doyle and Bishop, 1993) Su(tor)2-93 (Doyle and Bishop, 1993) Su(tor)2-113 (Doyle and Bishop, 1993) Su(tor)2-180 (Doyle and Bishop, 1993) Su(tor)2-187-1 (Doyle and Bishop, 1993) Su(tor)2-187-3 (Doyle and Bishop, 1993) Su(tor)2-191 (Doyle and Bishop, 1993) Su(tor)2-237 (Doyle and Bishop, 1993) Su(tor)2-278 (Doyle and Bishop, 1993) Su(tor)2-280 (Doyle and Bishop, 1993) Su(tor)2-321 (Doyle and Bishop, 1993) Su(tor)2-341 (Doyle and Bishop, 1993) Su(tor)2-399 (Doyle and Bishop, 1993) Su(tor)2-408 (Doyle and Bishop, 1993) Su(tor)3-1 (Doyle and Bishop, 1993) Su(tor)3-8 (Doyle and Bishop, 1993) Su(tor)3-91 (Doyle and Bishop, 1993) Su(tor)3-168 (Doyle and Bishop, 1993) Su(tor)3-293 (Doyle and Bishop, 1993) Su(tor)3-307 (Doyle and Bishop, 1993) Su(tor)3-333 (Doyle and Bishop, 1993) Su(tor)3-337 (Doyle and Bishop, 1993) Su(tor)3-344 (Doyle and Bishop, 1993) tor12D 14-3-3ε (Li and Li, 2003) 14-3-3ζ (Li and Li, 2003) bowl (Li and Li, 2003) Btk29A (Li et al., 2000) byn (Li and Li, 2003) csw (Li and Li, 2003) dpp (Li and Li, 2003) drk (Li and Li, 2003) Dsor1 (Li and Li, 2003) hkb (Li and Li, 2003) Hsp83 (Li and Li, 2003) lilli (Li and Li, 2003) Mad (Li and Li, 2003) phl (Li and Li, 2003) Ras85D (Li and Li, 2003) rl (Li and Li, 2003) Sos (Li and Li, 2003) Stat92E (Li et al., 2002, Li and Li, 2003) tkv (Li and Li, 2003) tll (Li and Li, 2003) tup (Li and Li, 2003) tor13D.hs.sev stumps (Vincent et al., 1998) tor13D dos (Luschnig et al., 2000) dpp (Strecker et al., 1991, Li and Li, 2003) gt (Strecker et al., 1991) Shc (Luschnig et al., 2000) torrv66 14-3-3ζ (Li et al., 1997) tsl (Furriols et al., 1998) torRX bcd (Schaeffer et al., 2000) Ras85D (Greenwood and Struhl, 1997) unspecified bcd (Werz et al., 2005) ems (Strecker et al., 1992) fog (Strecker et al., 1992) lin (Strecker et al., 1992) peb (Strecker et al., 1992) sog (Strecker et al., 1992) TfIIA-S (Zeidler et al., 1996) trk (Furriols et al., 1996) zen (Strecker et al., 1992)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 61589 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0003733 DroID - A comprehensive database of gene and protein interactions. • FBgn0003733 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 35717
Orthologs
Genome-wide drosophilid orthologs	Dana\GF11696 Dere\GG10716 Dgri\GH20010 Dmoj\GI19191 Dper\GL11534 Dpse\GA12603 Dsec\GM20762 Dvir\GJ20256 Dwil\GK19476 Dyak\GE23684
Curated drosophilid orthologs	Dpse\GA12603   Dsim\tor (Harr and Schlotterer, 2000)
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL010923-PA Aedes aegypti (mosquito) AGAP005763-PA Anopheles gambiae (mosquito) 86594 Branchiostoma floridae (lancelet) BGIBMGA003976-PA Bombyx mori (silk worm) ENSBTAP00000003980 Bos taurus (bovine) CN04163 Caenorhabditis brenneri (small nematode) CE12472 Caenorhabditis elegans (worm) ENSCINP00000019224 Ciona intestinalis (sea squirt) JA00003 Caenorhabditis japonica (nematode) CPIJ001844 Culex pipens (mosquito) RP47911 Caenorhabditis remanei (nematode) 175642 Hypsipyla robusta (cedar shoot borer) ISCW002017-PA Ixodes scapularis (deer tick) 119146 Lottia gigantea (owl limpet) 37485 Monocosta brevicollis (choanoflagellate) ENSMMUP00000011992 Macaca mulatta (rhesus monkey) XP_001607119.1 Nasonia vitripennis (wasp) PP08401 Pristionchus pacificus (nematode) 182742 Physcomitrella patens (moss) ENSPPYP00000011205 Pongo pygmaeus (orangutan) 82498 Phytophthora ramorum (sudden oak death pathogen) 771096 Populus trichocarpa (black cottonwood) 5056724 Sorghum bicolor (sorghum) NP_001034536.1 Tribolium castaneum (red flour beetle) ENSXETP00000020672 Xenopus tropicalis (frog) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • NV11573 FBgn0240844 FBgn0221474 FBgn0207396 FBgn0182001 FBgn0175643 FBgn0072649 FBgn0149143 FBgn0127474 FBgn0103021 FBgn0088736 CPIJ001844 BGIBMGA003976 ACYPI009483 AAEL010923 GLEAN_09906
Stocks & Reagents
Stocks Listed in FlyBase ( 25 )
Bloomington	17818 w1118; PBac{RB}tore00150 33627 y1 v1; P{TRiP.HMS00021}attP2
Harvard	- PBac{RB}tore00150
Kyoto	104475 w*; P{GawB}torNP3370 113244 w*; P{GawB}torNP3370
VDRC	v101154 P{KK105371}VIE-260B v1016 w1118; P{GD117}v1016 v4298 w1118; P{GD2613}v4298 v36280 w1118; P{GD14396}v36280 v4299 w1118; P{GD2613}v4299/TM3 v4300 w1118; P{GD2613}v4300/TM3 v36281 w1118; P{GD14396}v36281/TM3
	More stocks available...
Genomic Clones ( 2 )
	BACR04B09 BACR29D02 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 110 )
	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	RE49094
Other clones	DPiM_RE49094 IP17975 MIP23840 RT10495
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	LD04767 LD05421 LD06734 LD06992 LD07861 LD13289 LD14469 LD19571 LD19787 LD20457 LD20822 LD24992 LD31041 LD32428 LD44307 LD45474 LD45919 LD46792 RE17575 RE39619 RE41193 RE41301 RE47816 RE49057 RE52556 RE52748 RE63887
Other clones	37844 48508 69175 73184 193486 214283.36 214577.21 Dmel_dig_001265_0585_3622 Dmel_dig_004301_0660_2672 Dmel_dig_006725_0713_3350 Dmel_dig_009240_0680_3250 Dmel_dig_012427_0604_1080 EC053035 EC053051 EC055787 EK150027 EK172532 EK172850 EK196459 EK201943 EP10139 IP17675 IP17775 IP17875 RP004049556 RP004061231 RP004116663 RP004121194 RP004127452 RP004160694 RP004163453 RP004166353 RP004172864 RP004179655 RP004188837 RP004188976 RP004197342 RP004205547 RP004206941 RP004208076 RP004210430 RP004210445 RP004215943 RP004217536 RP004218290 RP004220439 RP004220520 RP004227710 RP004228200 RP004240234 RP004243878 RP004246618 RP004252963 RP004253268 RP004254041 RP004261564 RP004268800 RP004310285 RP004369861 RP004387496 RP004391179 RP004393002 RP004424656 RP004432575 RP004433047 RP004483224 RP004490861 RP004491984 RP004500215 RP004520637 RP004522071 RP004523648 RP004525331 RP004526762 RP004528640 RP004534057 RP004535191 RP004553087
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0003733 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 35717
Antibody Information
	polyclonal (Casanova and Struhl, 1989)
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of	Source for identity of: tor CG1389 (Gabrielian, 1999.11)
Source for database merge of
Additional comments
Other Comments
	When dsRNA constructs are made and transiently transfected into S2 cells in RNAi experiments, a decrease in the ratio of cells in prometaphase and metaphase versus the total number of mitotic cells seen. (Bettencourt-Dias et al., 2004) dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology. (Kiger et al., 2003) In tor gain-of-function embryos, primordial germ cells migrate out of the posterior midgut prematurely. They appear more abundant and motile than in wild-type embryos. (Li et al., 2003) ebi is required for Egfr but not tor-receptor-dependent gene expression. (Dong et al., 1999) Mutating all the tyrosine residues in the kinase insert results in a complete loss of function phenotype. Both phosphorylated tyrosine sites in the kinase domain activation loop are necessary for tor catalytic activity - mutation of these sites also leads to a complete tor loss of function phenotype. (Gayko et al., 1999) In a sample of 79 genes with multiple introns, 33 showed significant heterogeneity in G+C content among introns of the same gene and significant positive correspondence between the intron and the third codon position G+C content within genes. These results are consistent with selection adding against preferred codons at the start of genes. (Kliman and Eyre-Walker, 1998) Phylogenetic analysis of the PTK family. (Suga et al., 1997) phl Ser/Thr kinase activity is essential for its role in tor signal transduction. Altered forms of phl have distinct activity profiles indicating that each structural modification differentially affects the regulation and/or propagation of the tor signal by the mutant phl proteins. (Baek et al., 1996) tor does not affect the ability of bcd to bind DNA, but instead directs modification of bcd or of a potential bcd co-factor, which renders the bcd protein unable to activate transcription. (Bellaiche et al., 1996) Terminal structure development is regulated by the compensatory activities of positive and negative phosphotyrosine signaling sites on the tor receptor tyrosine kinase. (Cleghon et al., 1996) Different amounts of tor or trk molecules correlate with the expression of different zygotic genes, implicating changes in the number of activated tor molecules as one of the mechanisms defining differential gene expression. (Furriols et al., 1996) Nurse cell-specific genes are functional in the pseudonurse cells of otu mutants, but the transport of pum, otu, ovo and bcd RNAs to the cytoplasm is affected. (Heino et al., 1995) phl can be activated by tor in the complete absence of Ras85D, phl can be activated by RPTK in a Ras-independent pathway. (Hou et al., 1995) phl can be activated by tor in the complete absence of Ras85D function. (Hou et al., 1995) The tor response elements have been mapped in the tll promoter. The 11bp response element mediates repression of tll. This repression is lifted by activation of the tor pathway at the poles of the embryo. (Liaw et al., 1995) ksr functions in multiple receptor tyrosine kinase pathways. (Therrien et al., 1995) Ectopically activated tor acts as to silence the mus209 gene through its 5' homeodomain protein binding region. (Yamaguchi et al., 1995) 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) Distribution of tud protein in mutant embryos has been studied. Maternal genes such as bcd, tor and trk that are necessary for anteroposterior axis formation, but not required for germ cell formation or abdominal segmentation, have no effect on the distribution of tud protein. (Bardsley et al., 1993) The failure to accumulate tor protein at one or both of the poles leads to spatially inappropriate activity of more centrally located tor receptor: ectopic activity depends on the same gene functions normally required for activating tor. Ectopic activity reflects inappropriate diffusion of the ligand to more central regions of the body, and therefore concluded that the tor receptor not only transduces the spatial signal imparted by the tor ligand, but sequesters the ligand, ensuring its correct localization. (Casanova and Struhl, 1993) tll expression domain is restored in suppressed tor mutant embryos. (Doyle and Bishop, 1993) The role of tor in the regulation of run mRNA expression in the early embryo has been investigated. (Klingler and Gergen, 1993) Ecol\lacZ reporter gene constructs demonstrate the presence of tor maternal system cis-acting response elements in the 5' flanking region of tll. (Liaw and Lengyel, 1993) Heat shock induced expression, embryonic injection and female mosaic analysis demonstrates that Ras85D and its positive regulator Sos are involved in the tor signalling pathway. (Lu et al., 1993) In its anterior domain (labral primordia) cnc is activated by bicoid and torso maternal pathways. (Mohler, 1993) An artificial bcd responder gene composed of three bcd consensus binding sites driving Ecol\lacZ is activated by bcd and repressed by tor. This repression does not require tll or hkb. Phosphorylation resulting from the tor signal transduction pathway down-regulates transcriptional activation by the bcd morphogen. The normal phosphorylation changes that affect bcd during development do not occur in tor mutant embryos. (Ronchi et al., 1993) Biochemical analysis of the signal transduction pathway determining terminal structure development. (Sprenger et al., 1993) Portion of torso gene used in P element construct to provide activated phl protein, for study of phl and Ras85D functions in sevenless signal transduction pathway. (Dickson et al., 1992) csw functions downstream of tor. (Perkins et al., 1992) tor loss-of-function mutants delete the terminal regions in tll embryos, gain-of-function mutants expand the terminal domains. The maternal terminal system is necessary to activate tll expression in the terminal caps. (Pignoni et al., 1992) Injecting eggs with torso mRNA revealed that torso receptor tyrosine kinase activation is governed by an extracellular molecule produced at the terminal regions of the egg early in embryogenesis. When torso is absent this ligand fails to localise. Mutant ligand-binding torso proteins can suppress telson formation in a dominant negative manner indicating that the ligand is limited in amount. Analysis of torso mutations indicates that gain of function mutations causing ligand-independent activation map to the extracellular domain. (Sprenger and Nusslein-Volhard, 1992) The activation and spatial limitation of tll and hkb expression in the posterior region of the embryo is critically dependent on tor activity. (Bronner and Jaeckle, 1991) 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) Increased tor activity compensates for the absence of run activity to activate Sxl expression in the central and terminal regions. (Duffy and Gergen, 1991) tor has a repressive effect posteriorly and an inductive effect anteriorly on gt expression domain. (Eldon and Pirrotta, 1991) tor is responsible for specifying terminal structures. (Kraut and Levine, 1991) Mutations in maternal terminal class gene tor do not interact with RpII140wimp. (Parkhurst and Ish-Horowicz, 1991) tor plays a role in the specification of the anterior and posterior pole. (Ray et al., 1991) Zygotically active locus involved in the terminal developmental program in the embryo. (Strecker et al., 1991) The insertion of nos response elements (NREs) is sufficient to render maternal tor transcripts sensitive to repression by nos. (Wharton and Struhl, 1991) tor mutants exhibit deletion of the acron and telson. (Gaul and Jaeckle, 1990) Mature follicles are immunologically stained for asymmetric distribution of ecdysteroid-related antigen. During late oogenesis localisation of the antigen changes dramatically suggesting the antigen plays a role in early embryogenesis and, perhaps, in pattern formation. (Grau and Gutzeit, 1990) phl acts downstream of tor. (Ambrosio et al., 1989) tor protein is uniformly expressed along the surface membrane of early embryos, despite its localised activity at both poles. (Casanova and Struhl, 1989) Mutation in tor results in a maternal effect phenotype with defects during the early stages of gastrulation and defects in the anteroposterior axis. (Schupbach and Wieschaus, 1989) The product of the tor gene has a structure similar to receptor tyrosine kinase. (Sprenger et al., 1989) Involved in functions related to that of tll. (Strecker et al., 1988) Pattern elements from the anterior and posterior have been deleted in embryos of tor mutants. (Schupbach and Wieschaus, 1986) A single dose of tll1 from a tor11D/tor11D mother can partially rescue the tor11D mutant effect in the embryo (loss of abdominal segments); complete rescue may occur when the embryo is homozygous for tll1, receiving the gene from both parents (Strecker, Halsell, Fisher and Lipschitz, 1989). Injection of tor+ cytoplasm from early cleavage embryos can partially rescue tor loss-of-function mutants. ftz expression is reduced or lost in strong gain-of-function mutants (Klingerr et al., 1988; Strecker et al., 1989). phl mutations have been found to be epistatic over tor gain-of-function alleles (Nusslein-Volhard, Frohnhofer and Lehmann, 1987).   maternal-effect lethal embryos from homozygous mothers show alterations in the anterior-posterior pattern. Hypoactivity (loss-of-function) mutant embryos lack anteriormost head structures (labrum, dorsal bridges) as well as structures posterior to the seventh abdominal segment. Hyperactivity (gain of function) mutant embryos, on the other hand, show segment defects in the middle of the embryos, but may have enlarged terminal structures (Klingler, Erdelyi, Szabad and Nusslein-Volhard, 1988; Strecker, Halsell, Fisher and Lipschitz, 1989). A large number of revertants have been obtained from dominant or semidominant hypermorphic alleles. During cellularization at the blastoderm stage, hypoactivity mutant embryos show a 'pole hole' phenotype. A funnel of yolk-free cytoplasm with a small number of nuclei (between 10 and 20) is formed at the posterior pole, extending from the egg periphery to the inner yolk mass. At gastrulation the cephalic furrow is shifted toward the anterior and the germband extends all the way to the posterior end. Analysis of germ-line clones indicates that the torso mutant is germ-line autonomous (Schupbach and Wieschaus, 1986a).
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_057414 NM_001103741 NM_001103742 RefSeq (Proteins) • NP_476762 NP_001097211 NP_001097212 DDBJ / EMBL / GenBank • AAF59203 AAL49025 ABV53713 ABV53714 AC004283 AY071403 BI364057 CAA33247 CZ478045 X15150 Entrez Gene - A searchable database of RefSeq genes. • 35717 UniProtKB/Swiss-Prot • P18475
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Protein kinase, catalytic domain (IPR000719) Tyrosine-protein kinase, active site (IPR008266) Protein kinase-like domain (IPR011009) Fibronectin, type III (IPR003961) Protein kinase, ATP binding site (IPR017441) Tyrosine-protein kinase, receptor torso (IPR020724) Tyrosine-protein kinase, catalytic domain (IPR020635) Serine-threonine/tyrosine-protein kinase (IPR001245)
Linkouts
	BioGRID - A database of protein and genetic interactions • 61589 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0003733 DroID - A comprehensive database of gene and protein interactions. • FBgn0003733 DRSC - Results from RNAi screens. • FBgn0003733 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003733 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG1389-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0003733 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 35717 InParanoid A subset of ortholog calls from InParanoid. • AAEL010923-PA Aedes aegypti (mosquito) AGAP005763-PA Anopheles gambiae (mosquito) 86594 Branchiostoma floridae (lancelet) BGIBMGA003976-PA Bombyx mori (silk worm) ENSBTAP00000003980 Bos taurus (bovine) CN04163 Caenorhabditis brenneri (small nematode) CE12472 Caenorhabditis elegans (worm) ENSCINP00000019224 Ciona intestinalis (sea squirt) JA00003 Caenorhabditis japonica (nematode) CPIJ001844 Culex pipens (mosquito) RP47911 Caenorhabditis remanei (nematode) 175642 Hypsipyla robusta (cedar shoot borer) ISCW002017-PA Ixodes scapularis (deer tick) 119146 Lottia gigantea (owl limpet) 37485 Monocosta brevicollis (choanoflagellate) ENSMMUP00000011992 Macaca mulatta (rhesus monkey) XP_001607119.1 Nasonia vitripennis (wasp) PP08401 Pristionchus pacificus (nematode) 182742 Physcomitrella patens (moss) ENSPPYP00000011205 Pongo pygmaeus (orangutan) 82498 Phytophthora ramorum (sudden oak death pathogen) 771096 Populus trichocarpa (black cottonwood) 5056724 Sorghum bicolor (sorghum) NP_001034536.1 Tribolium castaneum (red flour beetle) ENSXETP00000020672 Xenopus tropicalis (frog) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /torstoll/torso1.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 35717 modMine - Data generated by the modENCODE project. • FBgn0003733 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • NV11573 FBgn0240844 FBgn0221474 FBgn0207396 FBgn0182001 FBgn0175643 FBgn0072649 FBgn0149143 FBgn0127474 FBgn0103021 FBgn0088736 CPIJ001844 BGIBMGA003976 ACYPI009483 AAEL010923 GLEAN_09906
Synonyms & Secondary IDs ( 8 )
Reported As
Symbol Synonym	CG1389 (Morrison et al., 2000, Keleman et al., 2009.8.5, Ni et al., 2010.12.1, Ni et al., 2011.7.19) splc (Schupbach and Wieschaus, 1989, ) Tor (Li et al., 2000, Hamilton et al., 2003, Fernandez-Minan et al., 2007, Bauer et al., 2010) tor (Friedman, 2007, McGregor, 2006, de las Heras and Casanova, 2006, Florence and Faller, 2008, Cinnamon et al., 2008, Baig-Lewis et al., 2007, Wang et al., 1994, Zeitlinger et al., 2007, Löhr et al., 2009, de Las Heras et al., 2009, Blanco and Gehring, 2008, Ding et al., 2010, Ventura et al., 2010, Sprenger et al., 1993, Grillo et al., 2011, Ajuria et al., 2011) TOR (Lee and Chung, 2007) Torso (Cinnamon et al., 2007, Kim et al., 2008)
Name Synonym	spliced   torso (Cui et al., 2008, Sprenger, 1994.12.21, Lai et al., 1995, Shishido et al., 1991, Coppey et al., 2007, Friedman, 2007, McGregor, 2006, De las Heras et al., 2007, Lemke et al., 2008, Petit et al., 2004, Florence and Faller, 2008, Xia et al., 2008, Zeitlinger et al., 2007, Rewitz et al., 2009, de Las Heras et al., 2009, Blanco and Gehring, 2008, Ding et al., 2010, Xiang et al., 2010, Sprenger et al., 1993, Ajuria et al., 2011) Torso (Wehrli and Baig-Lewis, 2006, Hariharan, 2005, Casanova, 2006, Ventura et al., 2008, Hamilton et al., 2003, Baig-Lewis et al., 2007, Fernandez-Minan et al., 2007, Coppey et al., 2008, Tanentzapf et al., 2006, Mao and Freeman, 2009, Berezhkovskii et al., 2009, Löhr et al., 2009, Ventura et al., 2010, Grillo et al., 2011)
Secondary FlyBase IDs
References ( 433 )
Generate a list of	All references relating to this gene ( 433 )
List References by type	Research paper  ( 252 ) Supplementary material  ( 6 ) Review  ( 108 ) Erratum  ( 1 ) Personal communication to FlyBase  ( 8 ) Abstract  ( 39 ) FlyBase analysis  ( 3 ) DNA/RNA sequence record  ( 1 ) Letter  ( 3 ) Stock list  ( 1 ) Curated genome annotation  ( 2 ) Protein sequence record  ( 1 ) Book  ( 1 ) Computer file  ( 4 ) Automatic genome annotation  ( 1 ) Spoof  ( 1 ) Conference report  ( 1 )
Recent research papers ( 7 )
	Ajuria et al., 2011, Development 138(5): 915--924 Capicua DNA-binding sites are general response elements for RTK signaling in Drosophila. [FBrf0212975] Gibbens et al., 2011, Development 138(13): 2693--2703 Neuroendocrine regulation of Drosophila metamorphosis requires TGF{beta}/Activin signaling. [FBrf0213899] Grillo et al., 2011, Genetics 187(2): 513--521 Control of Germline torso Expression by the BTB/POZ Domain Protein Pipsqueak Is Required for Embryonic Terminal Patterning in Drosophila. [FBrf0213010] Bauer et al., 2010, BMC Bioinformatics 11: 366 Dual-functioning transcription factors in the developmental gene network of Drosophila melanogaster. [FBrf0211408] Ding et al., 2010, Genetics 184(3): 717--729 Drosophila Raf's N terminus contains a novel conserved region and can contribute to torso RTK signaling. [FBrf0210411] Ventura et al., 2010, Dev. Biol. 344(1): 224--232 closca, a new gene required for both Torso RTK activation and vitelline membrane integrity. Germline proteins contribute to Drosophila eggshell composition. [FBrf0211365] Xiang et al., 2010, Dev. Biol. 344(2): 800--808 br regulates the expression of the ecdysone biosynthesis gene npc1. [FBrf0211381] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...