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General Information
Symbol
Dmel\tor
Species
D. melanogaster
Name
torso
Annotation Symbol
CG1389
Feature Type
FlyBase ID
FBgn0003733
Gene Model Status
Stock Availability
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]
Also Known As
splc
Key Links
Genomic Location
Cytogenetic map
Sequence location
2R:7,705,183..7,709,184 [-]
Recombination map
2-57
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the protein kinase superfamily. Tyr protein kinase family. (P18475)
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
Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
2

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
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0088938
3131
923
FBtr0339117
3116
918
Additional Transcript Data and Comments
Reported size (kB)
5.5, 3.6 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0088012
105.2
923
6.74
FBpp0308262
104.7
918
6.74
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)
Comments
External Data
Post Translational Modification
May be auto-phosphorylated on tyrosine residues.
(UniProt, P18475)
Linkouts
Sequences Consistent with the Gene Model
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
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR000719, InterPro:IPR017441
(assigned by InterPro )
inferred from biological aspect of ancestor with PANTHER:PTN002356460
(assigned by GO_Central )
Biological Process (13 terms)
Terms Based on Experimental Evidence (12 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:gcl; FB:FBgn0005695
inferred from genetic interaction with FLYBASE:Cul3; FB:FBgn0261268
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:trk; FB:FBgn0003751
inferred from genetic interaction with FLYBASE:Raf; FB:FBgn0003079
inferred from direct assay
inferred from expression pattern
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN002356460
(assigned by GO_Central )
Cellular Component (2 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN002356460
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN002356460
(assigned by GO_Central )
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
organism

Comment: maternally deposited

organism

Comment: rapidly degraded

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
radioisotope in situ
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
The Kr promoter directs expression of tor mRNA to the central portion of the embryo.
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.
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.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
tor protein is ubiquitously distributed over the cell surface.
The tor protein is expressed in the central portion of the embryo.
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.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
Reporter: P{GawB}torNP3370
Stage
Tissue/Position (including subcellular localization)
Reference
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
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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 9-10
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 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (19)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
3 of 15
Yes
No
 
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
 
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
 
2 of 15
No
Yes
2 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (19)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
3 of 15
Yes
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
Yes
2 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
Rattus norvegicus (Norway rat) (19)
2 of 13
Yes
No
2 of 13
Yes
No
2 of 13
Yes
No
2 of 13
Yes
No
2 of 13
Yes
No
2 of 13
Yes
No
2 of 13
Yes
No
2 of 13
Yes
No
2 of 13
Yes
No
2 of 13
Yes
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (17)
2 of 12
Yes
No
2 of 12
Yes
No
2 of 12
Yes
Yes
2 of 12
Yes
Yes
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
Yes
1 of 12
No
Yes
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
Danio rerio (Zebrafish) (27)
2 of 15
Yes
No
2 of 15
Yes
Yes
2 of 15
Yes
No
2 of 15
Yes
No
2 of 15
Yes
No
2 of 15
Yes
No
2 of 15
Yes
No
2 of 15
Yes
No
2 of 15
Yes
No
2 of 15
Yes
No
2 of 15
Yes
Yes
2 of 15
Yes
Yes
2 of 15
Yes
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (28)
3 of 15
Yes
Yes
3 of 15
Yes
Yes
3 of 15
Yes
Yes
3 of 15
Yes
Yes
3 of 15
Yes
Yes
3 of 15
Yes
Yes
3 of 15
Yes
Yes
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
Yes
2 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
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
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
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
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Culex quinquefasciatus
Southern house mosquito
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
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Tribolium castaneum
Red flour beetle
Rhodnius prolixus
Kissing bug
Acyrthosiphon pisum
Pea aphid
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
Ixodes scapularis
Black-legged tick
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
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (22)
4 of 10
3 of 10
3 of 10
3 of 10
3 of 10
2 of 10
2 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
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 )
    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.
    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
    Show neighbor-neighbor interactions:
    Select Layout:
    Legend:
    Protein
    RNA
    Selected Interactor(s)
    Interactions Browser

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    esyN Network Diagram
    esyN Network Key:
    Suppression
    Enhancement

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    suppressible
    suppressible
    suppressible
    suppressible
    suppressible
    suppressible
    suppressible
    suppressible
    suppressible
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    External Data
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    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.
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2R
    Recombination map
    2-57
    Cytogenetic map
    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)
    43E5-43E8
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (22)
    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
    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.

    cDNA Clones, End Sequenced (ESTs)
    RNAi and Array Information
    Linkouts
    DRSC - Results frm RNAi screens
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    Antibody Information
    Laboratory Generated Antibodies
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for identity of: tor CG1389
    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.
    dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.
    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.
    ebi is required for Egfr but not tor-receptor-dependent gene expression.
    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.
    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.
    Phylogenetic analysis of the PTK family.
    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.
    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.
    Terminal structure development is regulated by the compensatory activities of positive and negative phosphotyrosine signaling sites on the tor receptor tyrosine kinase.
    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.
    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.
    phl can be activated by tor in the complete absence of Ras85D, phl can be activated by RPTK in a Ras-independent pathway.
    phl can be activated by tor in the complete absence of Ras85D function.
    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.
    ksr functions in multiple receptor tyrosine kinase pathways.
    Ectopically activated tor acts as to silence the mus209 gene through its 5' homeodomain protein binding region.
    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.
    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.
    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.
    tll expression domain is restored in suppressed tor mutant embryos.
    The role of tor in the regulation of run mRNA expression in the early embryo has been investigated.
    Ecol\lacZ reporter gene constructs demonstrate the presence of tor maternal system cis-acting response elements in the 5' flanking region of tll.
    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.
    In its anterior domain (labral primordia) cnc is activated by bicoid and torso maternal pathways.
    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.
    Biochemical analysis of the signal transduction pathway determining terminal structure development.
    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.
    csw functions downstream of tor.
    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.
    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.
    The activation and spatial limitation of tll and hkb expression in the posterior region of the embryo is critically dependent on tor activity.
    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.
    Increased tor activity compensates for the absence of run activity to activate Sxl expression in the central and terminal regions.
    tor has a repressive effect posteriorly and an inductive effect anteriorly on gt expression domain.
    tor is responsible for specifying terminal structures.
    Mutations in maternal terminal class gene tor do not interact with RpII140wimp.
    tor plays a role in the specification of the anterior and posterior pole.
    Zygotically active locus involved in the terminal developmental program in the embryo.
    The insertion of nos response elements (NREs) is sufficient to render maternal tor transcripts sensitive to repression by nos.
    tor mutants exhibit deletion of the acron and telson.
    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.
    phl acts downstream of tor.
    tor protein is uniformly expressed along the surface membrane of early embryos, despite its localised activity at both poles.
    Mutation in tor results in a maternal effect phenotype with defects during the early stages of gastrulation and defects in the anteroposterior axis.
    The product of the tor gene has a structure similar to receptor tyrosine kinase.
    Involved in functions related to that of tll.
    Pattern elements from the anterior and posterior have been deleted in embryos of tor mutants.
    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).
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    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.
    GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
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    BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
    Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
    Flygut - An atlas of the Drosophila adult midgut
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