FlyBase Gene Report: 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

22 publicly available

Enzyme Name (EC)

Receptor protein-tyrosine kinase (2.7.10.1)

Gene Snapshot

torso (tor) encodes a receptor required for the specification of both terminal regions of the early embryo and for ecdysone synthesis in the prothoracic gland at pupariation. [Date last reviewed: 2019-03-14]

Other Summaries

Alliance
Auto summary
Pathway
Gene Group
UniProtKB
Red Book
Interactive Fly

Also Known As

splc

Key Links

Genomic Location

Cytogenetic map

Sequence location

2R:7,705,183..7,709,184 [-]

Recombination map

2-57

Sequence

Get Decorated FASTA

Genomic Maps

GBrowse JBrowse

Help me choose

Other Genome Views

The following external sites may use different assemblies or annotations than FlyBase.

Function

GO Summary Ribbons

[Detailed GO annotations]

Gene Group (FlyBase)

RECEPTOR TYROSINE KINASES

Pathway (FlyBase)

Torso Signaling Pathway Core Components

Protein Family (UniProt)

Belongs to the protein kinase superfamily. Tyr protein kinase family. (P18475)

Protein Signatures (InterPro)

Molecular Function (GO)
[Detailed GO annotations]

Experimental Evidence

protein binding; transmembrane receptor protein tyrosine kinase activity

Predictions / Assertions

ATP binding; transmembrane receptor protein tyrosine kinase activity

Catalytic Activity (EC)

Experimental Evidence

ATP + a [protein]-L-tyrosine = ADP + a [protein]-L-tyrosine phosphate (2.7.10.1)

Predictions / Assertions

ATP + a [protein]-L-tyrosine = ADP + a [protein]-L-tyrosine phosphate (2.7.10.1)

Summaries

Gene Group (FlyBase)

RECEPTOR TYROSINE KINASES -: Receptor tyrosine kinases (RTK) are single-pass transmembrane receptors expressed on the plasma membrane. Upon the binding of an extracellular signalling molecule (e.g. growth factors, hormones), RTKs dimerize leading to the activation of the intracellular tyrosine kinase domain and intermolecular phosphorylation. The phosphotyrosines function as specific sites for the assembly, phosphorylation and activation of downstream signaling molecules. (Adapted from PMID:20602996).

Pathway (FlyBase)

Torso Signaling Pathway Core Components -: The formation of Drosophila embryonic termini is controlled by the localized activation of Torso (tor) receptor tyrosine kinase. The Torso signaling pathway acts via the canonical Ras/Raf/MAP kinase cascade. (Adapted from FBrf0157176.)

Protein Function (UniProtKB)

Probable receptor tyrosine kinase which is required for determination of anterior and posterior terminal structures in the embryo (PubMed:2927509, PubMed:8423783). During postembryonic development, involved in the initiation of metamorphosis probably by inducing the production of ecdysone in response to prothoracicotropic hormone Ptth (PubMed:19965758). Binding to Ptth stimulates activation of canonical MAPK signaling leading to ERK phosphorylation (By similarity).

(UniProt, P18475)

Phenotypic Description (Red Book; Lindsley and Zimm 1992)

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).

Summary (Interactive Fly)

transmembrane - receptor tyrosine kinase - membrane receptor for Trunk - crucial for establishment of anterior and posterior cell identity of the embryo - required for ecdysone synthesis in the prothoracic gland at pupariation

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the GBrowse view of Dmel\tor or the JBrowse view of Dmel\tor for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)

Isoform displayed:

Pfam protein domains

InterPro name

classification

start

end

Protein Domains (via SMART)

Isoform displayed:

SMART protein domains

InterPro name

classification

start

end

Comments on Gene Model

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.49

Sequence Ontology: Class of Gene

nuclear_gene

protein_coding_gene

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

tor-RA

FBtr0088938

NM_057414

3131

923

tor-RD

FBtr0339117

NM_001103742

3116

918

Additional Transcript Data and 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

tor-PA

FBpp0088012

105.2

923

6.74

NP_476762

AAF59203

tor-PD

FBpp0308262

104.7

918

6.74

NP_001097212

ABV53714

Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments

Reported size (kDa)

923 (aa); 105 (kD predicted)

(Sprenger et al., 1989)

Comments

External Data

Post Translational Modification

May be auto-phosphorylated on tyrosine residues.

(UniProt, P18475)

Crossreferences

InterPro - A database of protein families, domains and functional sites

Linkouts

Sequences Consistent with the Gene Model

Nucleotide / Polypeptide Records

UniProt

P18475

Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\tor using the Feature Mapper tool.

External Data

Crossreferences

Linkouts

Gene Ontology (18 terms)

Molecular Function (3 terms)

Terms Based on Experimental Evidence (2 terms)

CV Term

Evidence

References

protein binding

inferred from physical interaction with FLYBASE:csw; FB:FBgn0000382

(Cleghon et al., 1998)

transmembrane receptor protein tyrosine kinase activity

inferred from direct assay

(Amarnath et al., 2017, Gayko et al., 1999, Cleghon et al., 1998, Cleghon et al., 1996, Sprenger et al., 1993)

inferred from mutant phenotype

(Sprenger and Nusslein-Volhard, 1992)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

ATP binding

inferred from electronic annotation with InterPro:IPR000719, InterPro:IPR017441

(assigned by InterPro )

(FlyBase Curators et al., 2004-)

transmembrane receptor protein tyrosine kinase activity

inferred from biological aspect of ancestor with PANTHER:PTN002356460

(assigned by GO_Central )

(Gaudet et al., 2011)

Biological Process (13 terms)

Terms Based on Experimental Evidence (12 terms)

CV Term

Evidence

References

chorion-containing eggshell pattern formation

inferred from high throughput mutant phenotype

(Schupbach and Wieschaus, 1991)

gastrulation

inferred from mutant phenotype

(Li et al., 2003)

metamorphosis

inferred from mutant phenotype

(Rewitz et al., 2009)

negative phototaxis

inferred from mutant phenotype

(Yamanaka et al., 2013)

negative regulation of multicellular organism growth

inferred from mutant phenotype

(Johnson et al., 2013)

peptidyl-tyrosine autophosphorylation

inferred from direct assay

(Sprenger et al., 1993)

inferred from mutant phenotype

(Gayko et al., 1999)

pole cell fate determination

inferred from genetic interaction with FLYBASE:gcl; FB:FBgn0005695

(Pae et al., 2017)

inferred from genetic interaction with FLYBASE:Cul3; FB:FBgn0261268

(Pae et al., 2017)

pole cell migration

inferred from mutant phenotype

(Li et al., 2003)

positive regulation of MAPK cascade

inferred from mutant phenotype

(Ghiglione et al., 1999)

positive regulation of Ras protein signal transduction

inferred from direct assay

(Amarnath et al., 2017)

terminal region determination

inferred from mutant phenotype

(Jimenez et al., 2000, Gayko et al., 1999, Cleghon et al., 1996, Furriols et al., 1996, Savant-Bhonsale and Montell, 1993, Sprenger and Nusslein-Volhard, 1992, Stevens et al., 1990, Casanova and Struhl, 1989, Sprenger et al., 1989)

torso signaling pathway

inferred from mutant phenotype

(Gayko et al., 1999, Cleghon et al., 1996, Furriols et al., 1996, Sprenger and Nusslein-Volhard, 1992)

inferred from genetic interaction with FLYBASE:trk; FB:FBgn0003751

(Casanova and Struhl, 1989)

inferred from genetic interaction with FLYBASE:tsl; FB:FBgn0003867

(Savant-Bhonsale and Montell, 1993, Stevens et al., 1990)

inferred from genetic interaction with FLYBASE:Raf; FB:FBgn0003079

(Ding et al., 2010)

inferred from direct assay

(Amarnath et al., 2017, Sprenger et al., 1993)

inferred from expression pattern

(Ghiglione et al., 1999)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

transmembrane receptor protein tyrosine kinase signaling pathway

inferred from biological aspect of ancestor with PANTHER:PTN002356460

(assigned by GO_Central )

(Gaudet et al., 2011)

Cellular Component (2 terms)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

integral component of plasma membrane

inferred from direct assay

(Amarnath et al., 2017, Pae et al., 2017)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

integral component of plasma membrane

inferred from biological aspect of ancestor with PANTHER:PTN002356460

(assigned by GO_Central )

(Gaudet et al., 2011)

receptor complex

inferred from biological aspect of ancestor with PANTHER:PTN002356460

(assigned by GO_Central )

(Gaudet et al., 2011)

Expression Data

Expression Summary Ribbons

Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.

For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.

Transcript Expression

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage

organism | medial

(Casanova and Struhl, 1993)

embryonic stage 1 -- 3

organism

Comment: maternally deposited

(Fisher et al., 2012)

embryonic stage 4 -- 6

organism

Comment: rapidly degraded

(Fisher et al., 2012)

embryonic stage 7 -- 8

pole cell

(Fisher et al., 2012)

embryonic stage 9 -- 10

germline cell

(Fisher et al., 2012)

embryonic stage 13 -- 16

germline cell

(Fisher et al., 2012)

embryonic stage 17

prothoracic gland

(Rewitz et al., 2009)

third instar larval stage

class IV dendritic arborizing neuron

(Yamanaka et al., 2013)

wandering third instar larval stage

prothoracic gland

(Rewitz et al., 2009)

brain | faint

(Rewitz et al., 2009)

northern blot

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 1 -- 9

organism

(Sprenger et al., 1989)

adult stage | female

(Sprenger et al., 1989)

radioisotope in situ

Stage

Tissue/Position (including subcellular localization)

Reference

cleavage stage

organism | ubiquitous

(Sprenger et al., 1989)

adult stage & oogenesis

nurse cell

(Sprenger et al., 1989)

germarium

(Sprenger et al., 1989)

adult stage oogenesis 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

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 ubiquitously distributed over the cell surface.

(Casanova and Struhl, 1993, Casanova and Struhl, 1989)

The tor protein is expressed in the central portion of the embryo.

(Casanova and Struhl, 1993)

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)

Marker for

Subcellular Localization

CV Term

Evidence

References

integral component of plasma membrane

inferred from direct assay

(Amarnath et al., 2017, Pae et al., 2017)

Expression Deduced from Reporters

Reporter: P{GawB}tor^NP3370

Stage

Tissue/Position (including subcellular localization)

Reference

third instar larval stage

class IV dendritic arborizing neuron

(Yamanaka et al., 2013)

High-Throughput Expression Data

Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\tor in GBrowse 2

RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region

View exonic expression by developmental stage for Dmel\tor

View exonic expression by tissue for Dmel\tor

Bulk Downloads

RNA-Seq RPKM values for all genes

Reference

See Gelbart and Emmert, 2013 for analysis details and data files for all genes.

Developmental Proteome: Life Cycle

Developmental Proteome: Embryogenesis

External Data and Images

Linkouts

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0003733

FLIGHT - Cell culture data for RNAi and other high-throughput technologies

FBgn0003733

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

CG1389-RA

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG1389

Flygut - An atlas of the Drosophila adult midgut

FBgn0003733

Images

FlyExpress - Embryonic expression images (BDGP data)

Show all FlyExpress processed images for Dmel\tor

Stages(s) 1-3
Stages(s) 4-6
Stages(s) 7-8
Stages(s) 9-10

FlyBase Wiki

Image Based Resources

Alleles, Insertions, and Transgenic Constructs

Classical and Insertion Alleles ( 94 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 33 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of tor

Transgenic constructs containing regulatory region of tor

Deletions and Duplications ( 43 )

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)Drl^rv3

(Heitzler et al., 1993)

Df(2R)Drl^rv5

(Heitzler et al., 1993)

Df(2R)Drl^rv15

(Heitzler et al., 1993)

Df(2R)Drl^rv17

(Heitzler et al., 1993)

Df(2R)Drl^rv20

(Heitzler et al., 1993)

Df(2R)Drl^rv21

(Heitzler et al., 1993)

Df(2R)Drl^rv22

(Heitzler et al., 1993)

Df(2R)Drl^rv25

(Heitzler et al., 1993)

Df(2R)Drl^rv28

(Heitzler et al., 1993)

Df(2R)Drl^rv30

(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)

Phenotypes

For more details about a specific phenotype click on the relevant allele symbol.

Lethality

Allele

lethal

lethal, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

tor^GD14396

lethal | cold sensitive

lethal | dominant | maternal effect | heat sensitive

tor^11D

lethal | gap | heat sensitive

tor^11D

lethal | heat sensitive, with Scer\GAL4^Bx-MS1096

tor^4021.UAS

lethal | maternal effect

tor^11D

lethal | recessive

tor⁴

lethal - all die before end of embryonic stage | germline clone | non-rescuable maternal effect | recessive

lethal - all die before end of embryonic stage | maternal effect | cold sensitive

tor^12D

lethal - all die before end of embryonic stage | maternal effect | recessive

lethal - all die before end of embryonic stage | maternal effect | recessive | heat sensitive

tor^11D

lethal - all die during embryonic stage, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

tor^GD14396

partially lethal - majority die

tor^Y630F

viable

tor^e00150

viable, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

viable, with Scer\GAL4^Mef2.PR

viable, with Scer\GAL4^pnr-MD237

viable, with Scer\GAL4^tin.CΔ4

Sterility

Allele

female semi-sterile, with Scer\GAL4^nos.PG, Scer\GAL4^VP16.nos.UTR, Scer\GAL4^VP16.otu

tor^GL00222

female sterile, with Scer\GAL4^{VP16.mat.αTub67C}

tor^HMS00021

female sterile (with Df(2R)Exel6054)

tor⁴

female sterile | dominant

tor^13D

female sterile | heat sensitive

tor^11D

female sterile | recessive

female sterile germline-dependent | dominant

tor^13D

fertile

tor^e00150

Other Phenotypes

Allele

decreased cell number | embryonic stage (with tor²)

tor^Deg

decreased cell number | embryonic stage (with tor^Deg)

tor²

developmental rate defective, with Scer\GAL4^phm.PO

tor^GD14396

developmental rate defective | dominant | heat sensitive

tor^11D

developmental rate defective | recessive

eclosion rhythm defective, with Scer\GAL4^phm.PO

tor^GD14396

eclosion rhythm defective, with Scer\GAL4^tim.PE

tor^GD14396

embryonic/larval segmentation phenotype | embryonic stage | maternal effect

tor^rv66

hyperactive | dominant

tor^13D

hypoactive | adult stage, with Scer\GAL4^elav.PU

tor^HMS00021

hypoactive | dominant

tor^13D

large body, with Scer\GAL4^phm.PO

tor^GD14396

large body | recessive

tor^unspecified

neurophysiology defective | third instar larval stage (with tor¹)

tor^e00150

neurophysiology defective | third instar larval stage (with tor^e00150)

tor¹

phototaxis defective | third instar larval stage, with Scer\GAL4^elav-C155

tor^GD14396

phototaxis defective | third instar larval stage, with Scer\GAL4^GMR.PFa

tor^GD14396

phototaxis defective | third instar larval stage, with Scer\GAL4^GMR.PFa, Scer\GAL4^ppk.1.9

tor^GD14396

phototaxis defective | third instar larval stage, with Scer\GAL4^Kr.5.1

tor^GD14396

phototaxis defective | third instar larval stage, with Scer\GAL4^ppk.1.9

tor^GD14396

phototaxis defective | third instar larval stage, with Scer\GAL4^αTub84B.PL

tor^GD14396

phototaxis defective | third instar larval stage (with tor¹)

tor^e00150

phototaxis defective | third instar larval stage (with tor^e00150)

tor¹

size defective

tor^rv66

size defective | larval stage | somatic clone, with Scer\GAL4^NP0394

tor^GD14396

small body | dominant | male limited | heat sensitive

tor^11D

some die during embryonic stage | maternal effect | cold sensitive

tor^12D

some die during embryonic stage | non-rescuable maternal effect | recessive

some die during embryonic stage | recessive | maternal effect | recessive

some die during embryonic stage | recessive | maternal effect | recessive | heat sensitive

tor^11D

terminal phenotype | germline clone

tor^11D

terminal phenotype | germline clone | maternal effect

visible, with Scer\GAL4^Bx-MS1096

tor^4021.UAS

visible | heat sensitive

tor^11D

wild-type

Phenotype manifest in

Allele

abdomen | posterior

tor^rv66

abdominal 8 ventral denticle belt | maternal effect

tor^rv66

abdominal 11 anal tuft | embryonic stage

tor^rv66

abdominal segment

tor^11D

abdominal segment 8 | embryonic stage | maternal effect

tor^rv66

abdominal segment 9

abdominal segment 10

abdominal segment 11

tor⁴

adult labral sense organ

tor⁴

anterior midgut primordium | maternal effect | heat sensitive

tor^11D

cephalic furrow

cephalic furrow | maternal effect

cephalic furrow | maternal effect | heat sensitive

tor^11D

cephalopharyngeal skeleton

tor¹

cephalopharyngeal skeleton | maternal effect | cold sensitive

tor^12D

denticle belt

tor^12D

denticle belt | maternal effect

tor^12D

denticle belt | maternal effect | cold sensitive

tor^12D

denticle belt | maternal effect | heat sensitive

tor^11D

dorsal bridge

tor^unspecified

dorsopharyngeal organ

tor⁴

embryo

embryo | extended germ band stage | maternal effect

embryo | gastrula stage

tor^rv66

embryo | germline clone | maternal effect

embryo | maternal effect, with Scer\GAL4^nos.PG, Scer\GAL4^VP16.nos.UTR, Scer\GAL4^VP16.otu

tor^GL00222

embryo | maternal effect, with Scer\GAL4^{VP16.mat.αTub67C}

tor^HMS00021

embryo | posterior

embryonic/first instar larval cuticle

embryonic/first instar larval cuticle | maternal effect

embryonic/first instar larval cuticle | maternal effect (with tor^12D)

tor^rv66

embryonic/first instar larval cuticle | maternal effect (with tor^rv66)

tor^12D

embryonic/first instar larval cuticle | maternal effect | cold sensitive

tor^12D

embryonic/first instar larval cuticle | maternal effect | heat sensitive

tor^11D

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

tor^rv66

embryonic/larval optic lobe | embryonic stage

tor⁴

embryonic/larval posterior spiracle | embryonic stage

tor¹

embryonic abdominal segment 2 | heat sensitive

tor^11D

embryonic abdominal segment 3 | heat sensitive

tor^11D

embryonic abdominal segment 4 | heat sensitive

tor^11D

embryonic abdominal segment 4 | maternal effect

tor^11D

embryonic abdominal segment 5 | heat sensitive

tor^11D

embryonic abdominal segment 7

tor^unspecified

embryonic abdominal segment 8

embryonic abdominal segment | gap | heat sensitive

tor^11D

embryonic abdominal segment | maternal effect

tor^12D

embryonic abdominal segment | maternal effect | heat sensitive

tor^11D

embryonic anal pad

tor^rv66

embryonic epidermis

embryonic head

embryonic head | heat sensitive

tor^11D

embryonic head | maternal effect | posterior | heat sensitive

tor^11D

embryonic labial segment

tor^unspecified

embryonic labral segment | maternal effect

embryonic telson

tor^rv66

embryonic thoracic segment | gap | heat sensitive

tor^11D

embryonic thoracic segment | maternal effect | heat sensitive

tor^11D

endoderm

tor^13D

filzkorper

filzkorper | ectopic | maternal effect

tor^12D

filzkorper | embryonic stage

tor^rv66

filzkorper | embryonic stage | maternal effect

tor^rv66

filzkorper | maternal effect | cold sensitive

tor^12D

germline cell

tor^+t11.5

head

tor⁴

labral segment

tor⁴

labrum

labrum | embryonic stage

tor^rv66

nucleus | larval stage | somatic clone, with Scer\GAL4^NP0394

tor^GD14396

ocular segment

pharyngeal monoscolopidial chordotonal organ

tor⁴

pole cell | ectopic | maternal effect

tor^rv66

pole cell | embryonic stage (with tor²)

tor^Deg

pole cell | embryonic stage (with tor^Deg)

tor²

posterior midgut primordium

tor^rv66

posterior midgut primordium | maternal effect

prothoracic gland | larval stage | somatic clone, with Scer\GAL4^NP0394

tor^GD14396

pupa, with Scer\GAL4^phm.PO

tor^GD14396

salivary gland common duct primordium | maternal effect

tor^rv66

salivary gland common duct primordium | maternal effect | heat sensitive

tor^11D

stomatogastric nervous system

tor⁴

stomatogastric nervous system | embryonic stage

tor⁴

telson

telson | embryonic stage | maternal effect

tor^rv66

telson | maternal effect

ventral furrow

ventral furrow | maternal effect | heat sensitive

tor^11D

visceral mesoderm

tor^13D

wing, with Scer\GAL4^Bx-MS1096

tor^4021.UAS

Orthologs

Downloads

Download All DIOPT Orthologs

Download All OrthoDB Orthologs

Human Orthologs (via DIOPT v7.1)

Homo sapiens (Human) (19)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Hsap\PDGFRB

3 of 15

Yes

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

Yes

Hsap\TEK

2 of 15

Yes

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

Yes

Hsap\TADA2B

1 of 15

Hsap\TIE1

1 of 15

Model Organism Orthologs (via DIOPT v7.1)

Mus musculus (laboratory mouse) (19)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Mmus\Pdgfrb

3 of 15

Yes

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

Yes

Mmus\Tek

2 of 15

Yes

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

Yes

Mmus\Tada2b

1 of 15

Mmus\Tie1

1 of 15

Rattus norvegicus (Norway rat) (19)

Rnor\Csf1r

2 of 13

Yes

Rnor\Fgfr1

2 of 13

Yes

Rnor\Fgfr2

2 of 13

Yes

Rnor\Fgfr3

2 of 13

Yes

Rnor\Fgfr4

2 of 13

Yes

Rnor\Kit

2 of 13

Yes

Rnor\Pdgfra

2 of 13

Yes

Rnor\Pdgfrb

2 of 13

Yes

Rnor\Ret

2 of 13

Yes

Rnor\Styk1

2 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Yes

Rnor\Tada2b

1 of 13

Rnor\Tek

1 of 13

Rnor\Tie1

1 of 13

Xenopus tropicalis (Western clawed frog) (17)

Xtro\fgfr2

2 of 12

Yes

Xtro\fgfr3

2 of 12

Yes

Xtro\styk1

2 of 12

Yes

Xtro\tek

2 of 12

Yes

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

Yes

Xtro\pdgfrl

1 of 12

Yes

Xtro\ret

1 of 12

Xtro\tada2b

1 of 12

Xtro\tie1

1 of 12

Danio rerio (Zebrafish) (27)

Drer\csf1ra

2 of 15

Yes

Drer\csf1rb

2 of 15

Yes

Drer\fgfr1a

2 of 15

Yes

Drer\fgfr2

2 of 15

Yes

Drer\flt3

2 of 15

Yes

Drer\kita

2 of 15

Yes

Drer\kitb

2 of 15

Yes

Drer\pdgfra

2 of 15

Yes

Drer\pdgfrb

2 of 15

Yes

Drer\ret

2 of 15

Yes

Drer\si:ch73-206d17.1

2 of 15

Yes

Drer\si:dkey-14o18.6

2 of 15

Yes

Drer\styk1

2 of 15

Yes

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

Yes

Drer\si:ch211-167j9.5

1 of 15

Yes

Drer\tek

1 of 15

Drer\tie1

1 of 15

Caenorhabditis elegans (Nematode, roundworm) (28)

Cele\kin-15

3 of 15

Yes

Cele\kin-23

3 of 15

Yes

Cele\kin-30

3 of 15

Yes

Cele\old-1

3 of 15

Yes

Cele\old-2

3 of 15

Yes

Cele\R09D1.12

3 of 15

Yes

Cele\R151.1

3 of 15

Yes

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

2 of 15

Yes

Cele\ver-4

2 of 15

Cele\B0252.1

1 of 15

Cele\F40G9.8

1 of 15

Yes

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

1 of 15

Yes

Cele\ver-1

1 of 15

Cele\ver-3

1 of 15

Cele\W03A5.1

1 of 15

Yes

Cele\W04G5.10

1 of 15

Cele\Y38H6C.20

1 of 15

Cele\Y50D4B.6

1 of 15

Arabidopsis thaliana (thale-cress) (0)

No records found.

Saccharomyces cerevisiae (Brewer's yeast) (0)

No records found.

Schizosaccharomyces pombe (Fission yeast) (0)

No records found.

Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091901VB )

Organism

Common Name

Gene

AAA Syntenic Ortholog

Multiple Dmel Genes in this Orthologous Group

Drosophila melanogaster

fruit fly

Dmel\tor

Drosophila suzukii

Spotted wing Drosophila

Drosophila simulans

Drosophila sechellia

Drosophila erecta

Drosophila yakuba

Drosophila ananassae

Drosophila pseudoobscura pseudoobscura

Dpse\GA12603

Drosophila persimilis

Dper\GL11534

Drosophila willistoni

Dwil\GK19476

Drosophila virilis

Dvir\GJ20256

Drosophila mojavensis

Dmoj\GI19191

Drosophila grimshawi

Dgri\GH20010

Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG0915019N )

Organism

Common Name

Gene

Multiple Dmel Genes in this Orthologous Group

Musca domestica

House fly

Mdoa\17017455

Musca domestica

House fly

Mdoa\17019712

Glossina morsitans

Tsetse fly

Gmor\GMOY010892

Aedes aegypti

Yellow fever mosquito

Aaeg\AAEL010923

Aedes aegypti

Yellow fever mosquito

Aaeg\AAEL002404

Anopheles darlingi

American malaria mosquito

Adar\ADAC010295

Anopheles gambiae

Malaria mosquito

Agam\AGAP005763

Culex quinquefasciatus

Southern house mosquito

Cqui\CPIJ000058

Culex quinquefasciatus

Southern house mosquito

Cqui\CPIJ001844

Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W09KD )

Organism

Common Name

Gene

Multiple Dmel Genes in this Orthologous Group

Heliconius melpomene

Postman butterfly

Hmel\HMEL006361

Linepithema humile

Argentine ant

Lhum\LH12414

Megachile rotundata

Alfalfa leafcutting bee

Mrot\MROTU:001c0b

Nasonia vitripennis

Parasitic wasp

Nvit\Nasvi2EG003555

Tribolium castaneum

Red flour beetle

Tcas\tor

Rhodnius prolixus

Kissing bug

Rpro\RPRC001078

Acyrthosiphon pisum

Pea aphid

Apim\ACYPI009483

Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X09GE )

Organism

Common Name

Gene

Multiple Dmel Genes in this Orthologous Group

Strigamia maritima

European centipede

Smar\SMAR012984

Ixodes scapularis

Black-legged tick

Isca\ISCW007406

Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0CQZ )

Organism

Common Name

Gene

Multiple Dmel Genes in this Orthologous Group

Strongylocentrotus purpuratus

Purple sea urchin

Spur\Sp-Tk9

Strongylocentrotus purpuratus

Purple sea urchin

Spur\Sp-Tada2b

Strongylocentrotus purpuratus

Purple sea urchin

Spur\SPU_003252gn

Strongylocentrotus purpuratus

Purple sea urchin

Spur\Sp-Ret

Strongylocentrotus purpuratus

Purple sea urchin

Spur\Sp-Fgfr

Strongylocentrotus purpuratus

Purple sea urchin

Spur\Sp-Tk8

Strongylocentrotus purpuratus

Purple sea urchin

Spur\Sp-Cret

Strongylocentrotus purpuratus

Purple sea urchin

Spur\Sp-Cret_1

Ciona intestinalis

Vase tunicate

Cint\ENSCING00000001931

Ciona intestinalis

Vase tunicate

Cint\fgfr

Gallus gallus

Domestic chicken

Ggal\RET

Gallus gallus

Domestic chicken

Ggal\FGFR4

Gallus gallus

Domestic chicken

Ggal\FGFR1

Gallus gallus

Domestic chicken

Ggal\FGFR3

Gallus gallus

Domestic chicken

Ggal\TADA2B

Gallus gallus

Domestic chicken

Ggal\FGFR2

Paralogs

Downloads

Download All DIOPT Paralogs

Paralogs (via DIOPT v7.1)

Drosophila melanogaster (Fruit fly) (22)

Gene Symbol

Score

Source

Alignment

4 of 10

btl

3 of 10

Cad96Ca

3 of 10

htl

3 of 10

Tie

3 of 10

Pvr

2 of 10

Ret

2 of 10

Ada2b

1 of 10

Alk

1 of 10

CG10702

1 of 10

Ddr

1 of 10

dnt

1 of 10

Drl-2

1 of 10

drl

1 of 10

drpr

1 of 10

Egfr

1 of 10

Eph

1 of 10

InR

1 of 10

Nrk

1 of 10

Ror

1 of 10

sev

1 of 10

Wsck

1 of 10

Human Disease Associations

FlyBase Human Disease Model Reports

Disease Model Summary Ribbon

Disease Ontology (DO) Annotations

Models Based on Experimental Evidence ( 0 )

Allele

Disease

Evidence

References

Potential Models Based on Orthology ( 2 )

Human Ortholog

Disease

Evidence

References

PDGFRB; platelet derived growth factor receptor beta

model of basal ganglia calcification

IEA

(FlyBase, 2019-)

model of infantile myofibromatosis

IEA

(FlyBase, 2019-)

Modifiers Based on Experimental Evidence ( 0 )

Allele

Disease

Interaction

References

Comments on Models/Modifiers Based on Experimental Evidence ( 0 )

Disease Associations of Human Orthologs (via DIOPT v7.1 and OMIM)

Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.

Homo sapiens (Human)

Gene name

Score

OMIM

OMIM Phenotype

DO term

Complementation?

Transgene?

PDGFRB; platelet derived growth factor receptor beta

3 of 15

173410

Functional Complementation Data

Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.

Interactions

Summary of Physical Interactions

esyN Network Diagram

Interactions Browser

View in Interactions Browser

Please see the Physical Interaction reports below for full details

protein-protein

Physical Interaction

Assay

References

tor

bimolecular fluorescence complementation, fluorescence microscopy

(Amarnath et al., 2017)

tor - gcl

affinity technology, Identification by mass spectrometry, anti tag coimmunoprecipitation, anti tag western blot, western blot

(Pae et al., 2017)

Summary of Genetic Interactions

esyN Network Diagram

esyN Network Key:

Suppression

Enhancement

View in Interactions Browser

Please look at the allele data for full details of the genetic interactions

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

suppressible

(Li and Li, 2003)

suppressible

(Li and Li, 2003, Li et al., 1997)

tor

suppressible

bcd

(Schaeffer et al., 2000)

tor

suppressible

bowl

(Li and Li, 2003, Doyle and Bishop, 1993)

suppressible

(Li et al., 2000)

suppressible

(Li and Li, 2003)

enhanceable

(de las Heras and Casanova, 2006)

suppressible

(Li and Li, 2003)

suppressible

(Luschnig et al., 2000)

tor

enhanceable

dpp

(Li and Li, 2003, Strecker et al., 1991)

suppressible

(Li and Li, 2003)

suppressible

(Li and Li, 2003, Doyle and Bishop, 1993)

tor

suppressible

Dsor1

(Li and Li, 2003, Lim et al., 1997)

tor

enhanceable

fog

(Johnson et al., 2017)

tor

suppressible

(Strecker et al., 1991)

suppressible

(Li and Li, 2003)

suppressible

(Li and Li, 2003)

suppressible

(Li and Li, 2003)

suppressible

(Li and Li, 2003)

enhanceable

(Ding et al., 2010)

suppressible

(Li and Li, 2003)

suppressible

(Rewitz et al., 2009, Li and Li, 2003, Doyle and Bishop, 1993)

tor

suppressible

(Li and Li, 2003, Brunner et al., 1994)

tor

suppressible

Shc

(Luschnig et al., 2000)

tor

suppressible

Sos

(Li and Li, 2003, Doyle and Bishop, 1993)

cic|tor|trk

suppressible

Stat92E

(de las Heras and Casanova, 2006)

tor

suppressible

Stat92E

(Li and Li, 2003, Li et al., 2002)

tor

suppressible

sty

(Casci et al., 1999)

tor

suppressible

Su(tor)1-1

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)1-106

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)1-163

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-3

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-4

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-15

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-48

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-56

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-87

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-93

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-113

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-180

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-187-1

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-187-3

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-191

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-237

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-278

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-280

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-321

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-341

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-399

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)2-408

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)3-1

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)3-8

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)3-91

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)3-168

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)3-293

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)3-307

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)3-333

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)3-337

(Doyle and Bishop, 1993)

tor

suppressible

Su(tor)3-344

(Doyle and Bishop, 1993)

suppressible

(Li and Li, 2003)

suppressible

(Li and Li, 2003)

suppressible

(Johnson et al., 2013)

tor

enhanceable

tsl

(Johnson et al., 2013)

tor

suppressible

tup

(Li and Li, 2003)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

bcd

enhanceable

tor

(Werz et al., 2005)

cic

enhanceable

tor|trk

(de las Heras and Casanova, 2006)

clos

suppressible

tor

(Ventura et al., 2010)

fog

suppressible

tor

(Johnson et al., 2017)

suppressible

(Pae et al., 2017)

suppressible

(Vallejo et al., 2015)

psq

suppressible

tor

(Grillo et al., 2011)

stumps

suppressible

tor

(Vincent et al., 1998)

tsl

suppressible

tor

(Furriols et al., 1998)

tsl

enhanceable

tor

(Johnson et al., 2013)

External Data

Linkouts

BioGRID - A database of protein and genetic interactions.

61589

DroID - A comprehensive database of gene and protein interactions.

FBgn0003733

InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.

35717

MIST (genetic) - An integrated Molecular Interaction Database

MIST (protein-protein) - An integrated Molecular Interaction Database

Pathways

Gene Group - Pathway Membership (FlyBase)

Torso Signaling Pathway Core Components -: The formation of Drosophila embryonic termini is controlled by the localized activation of Torso (tor) receptor tyrosine kinase. The Torso signaling pathway acts via the canonical Ras/Raf/MAP kinase cascade. (Adapted from FBrf0157176.)

External Data

Linkouts

KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

P18475

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

2-57

Cytogenetic map

43E11-43E12

Sequence location

2R:7,705,183..7,709,184 [-]

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

(FlyBase, 2016, Schupbach and Wieschaus, 1989, Tearle and Nusslein-Volhard, 1987)

2-56.9

(Comeron et al., 2012)

Left of (cM)

(Szabad et al., 1989)

(Tearle and Nusslein-Volhard, 1987)

Right of (cM)

(Szabad et al., 1989)

Notes

Stocks and Reagents

Stocks (22)

Bloomington

17818

w¹¹¹⁸; PBac{RB}tor^e00150

33627

y¹ v¹; P{TRiP.HMS00021}attP2

58312

y¹ v¹; P{TRiP.HMJ22419}attP40

Harvard

e00150

PBac{RB}tor^e00150

Kyoto

104475

w^*; P{GawB}tor^NP3370

113244

w^*; P{GawB}tor^NP3370

VDRC

v101154

P{KK105371}VIE-260B

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 (11)

Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

cDNA Clones (99)

Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see GBrowse for alignment of the cDNAs and ESTs to the gene model.

cDNA clones, fully sequences

BDGP DGC clones

RE49094

Other clones

Drosophila Genomics Resource Center cDNA clones

For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

FBgn0003733

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0003733

GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents

35717

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Casanova and Struhl, 1989)

Commercially Available Antibodies

Other Information

Relationship to Other Genes

Source for database identify 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 Jäckle, 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 RpII140^wimp.

(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 Jäckle, 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 tor^11D/tor^11D mother can partially rescue the tor^11D 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).

Origin and Etymology

Discoverer

Etymology

Identification

External Crossreferences and Linkouts ( 45 )

Sequence Crossreferences

NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.

35717

GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.

GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.

RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.

UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information

P18475

Other crossreferences

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0003733

Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones

FBgn0003733

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG1389

Flygut - An atlas of the Drosophila adult midgut

FBgn0003733

GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents

35717

iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)

TC009906

InterPro - A database of protein families, domains and functional sites

KEGG Genes - Molecular building blocks of life in the genomic space.

Dmel_CG1389

KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.

MAPK

modMine - A data warehouse for the modENCODE project

FBgn0003733

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

P18475

Linkouts

BioGRID - A database of protein and genetic interactions.

61589

DPiM - Drosophila Protein interaction map

FBgn0003733

DroID - A comprehensive database of gene and protein interactions.

FBgn0003733

DRSC - Results frm 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 - An integrated database for Drosophila genomics

FBgn0003733

Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution

Interactive Fly

InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.

35717

KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.

MAPK signaling pathway - fly

MIST (genetic) - An integrated Molecular Interaction Database

MIST (protein-protein) - An integrated Molecular Interaction Database

Synonyms and Secondary IDs (9)

Reported As

Symbol Synonym

CG1389

(Yu et al., 2016, McEachern et al., 2014, Vogel et al., 2013, Ni et al., 2011.7.19, Nässel and Winther, 2010, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Morrison et al., 2000)

TOR

(Álvarez-Rendón et al., 2018, Lee et al., 2018, Schultzhaus et al., 2018, Kapahi et al., 2017, Aradhya et al., 2015, Wang et al., 2014, Whitaker et al., 2014, Erkosar et al., 2013, Jenkins et al., 2013, Homem and Knoblich, 2012, Mirth and Shingleton, 2012, Alic et al., 2011, Harrison and Haddad, 2011, Baudot et al., 2008, Lee and Chung, 2007)

Tor

(Richardson and Portela, 2018, Amarnath et al., 2017, Goyal et al., 2017, Goyal et al., 2017, Di Cara et al., 2015, Buchon et al., 2013, Johnson et al., 2013, Jaeger et al., 2012, Klusza and Deng, 2011, Bauer et al., 2010, Fernández-Miñán et al., 2007, Hamilton et al., 2003, Li et al., 2000)

Torso

(Pae et al., 2017, Danielsen et al., 2014, Hombría and Sotillos, 2013, Mirth and Shingleton, 2012, Gehring, 2011, Stec and Zeidler, 2011, Tsurumi et al., 2011, Kim et al., 2008, Cinnamon et al., 2007)

dTor

(Bednářová et al., 2018)

splc

(Schupbach and Wieschaus, 1989)

tor

(Myat et al., 2019, Skelly et al., 2019, Aboudhiaf et al., 2018, Chiku et al., 2018, Singh and Chowdhuri, 2018, Christesen et al., 2017, Liaw, 2016, Choi and Aquadro, 2015, Hsu et al., 2015, Parker and Struhl, 2015, Vallejo et al., 2015, Boisclair Lachance et al., 2014, Henstridge et al., 2014, Sopko et al., 2014, Villa-Cuesta et al., 2014, Johnson et al., 2013, Ozkan et al., 2013, Sopko and Perrimon, 2013, Chen et al., 2012, Grillo et al., 2012, Grimm et al., 2012, Meier et al., 2012, Ajuria et al., 2011, Grillo et al., 2011, Papatsenko and Levine, 2011, Ding et al., 2010, Ventura et al., 2010, Chen et al., 2009, de Las Heras et al., 2009, Löhr et al., 2009, Zhou et al., 2009, Blanco and Gehring, 2008, Cinnamon et al., 2008, Florence and Faller, 2008, Baig-Lewis et al., 2007, Friedman and Perrimon, 2007, Zeitlinger et al., 2007, de las Heras and Casanova, 2006, McGregor, 2006, Wratten et al., 2006, Wang et al., 1994, Sprenger et al., 1993)

torso

(Di Cara and King-Jones, 2016, Rahimi et al., 2016)

Name Synonyms

spliced

torso

(Shimell et al., 2018, Moeller et al., 2017, Selcho et al., 2017, Eichhorn et al., 2016, Jun et al., 2016, Niwa and Niwa, 2016, Ou et al., 2016, Komura-Kawa et al., 2015, Wang et al., 2015, Cantera et al., 2014, Enya et al., 2014, Johnson et al., 2013, Yamanaka et al., 2013, Miller et al., 2012, Ajuria et al., 2011, Ou et al., 2011, Tsurumi et al., 2011, Ding et al., 2010, Xiang et al., 2010, de Las Heras et al., 2009, Rewitz et al., 2009, Blanco and Gehring, 2008, Cui et al., 2008, Florence and Faller, 2008, Ishihara and Shibata, 2008, Lemke et al., 2008, Xia et al., 2008, Coppey et al., 2007, De las Heras et al., 2007, Friedman and Perrimon, 2007, Zeitlinger et al., 2007, McGregor, 2006, Petit et al., 2004, Lai et al., 1995, Sprenger, 1994.12.21, Sprenger et al., 1993, Shishido et al., 1991)

Secondary FlyBase IDs

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