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General Information
Symbol
Dmel\Tl
Species
D. melanogaster
Name
Toll
Annotation Symbol
CG5490
Feature Type
FlyBase ID
FBgn0262473
Gene Model Status
Stock Availability
Gene Snapshot
Toll (Tl) encodes a transmembrane receptor that activates the Tl intracellular signaling pathway upon binding the ligand encoded by spz. It is involved in dorso-ventral embryonic patterning and immunity. [Date last reviewed: 2019-03-14]
Also Known As
Toll-1, EP1051, T1, dToll
Key Links
Genomic Location
Cytogenetic map
Sequence location
3R:26,799,041..26,842,403 [+]
Recombination map
3-92
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 Toll-like receptor family. (P08953)
Summaries
Gene Group (FlyBase)
DORSAL GROUP -
The Dorsal group genes encode components of the Toll pathway and were initially identified by maternal effect mutations resulting in the dorsalization of the embryo. (Adapted from FBrf0091014 and FBrf0223077).
TOLL RECEPTORS -
The Toll/Toll-like receptor (TLR) family members are type-I single-pass transmembrane receptors with an intracellular Toll IL-1R homology (TIR) domain and extracellular Leucine-rich repeats. The archetypal family member, Tl, was first characterized for its role in embryonic dorsal/ventral patterning. In larval and adult stages, Tl is involved with innate immune response. Tl is activated by binding an endogenous spatzle ligand and activation of NF-κB is the target of canonical Toll/Toll-like receptor signaling. Non-canonical signaling pathways are less well characterized. (Adapted from FBrf0130108, FBrf0212564 and FBrf0239503).
Pathway (FlyBase)
Toll-NF-KappaB Signaling Pathway Core Components -
In Drosophila, the canonical Toll signaling pathway is initiated by the binding of a spatzle ligand to Toll (Tl) or a Toll-like receptor leading to the nuclear localization of the NF-κB (dl or Dif) transcription factor. (Adapted from FBrf0091014 and FBrf0223077).
Protein Function (UniProtKB)
Receptor for the cleaved activated form of spz, spaetzle C-106 (PubMed:12872120). Binding to spaetzle C-106 activates the Toll signaling pathway and induces expression of the antifungal peptide drosomycin (PubMed:12872120, PubMed:8808632, PubMed:10973475). Component of the extracellular signaling pathway that establishes dorsal-ventral polarity in the embryo (PubMed:3931919). Promotes heterophilic cellular adhesion (PubMed:2124970). Involved in synaptic targeting of motoneurons RP5 and V to muscle 12 (M12); functions as a repulsive cue inhibiting motoneuron synapse formation on muscle 13 (M13) to guide RP5 and V to the neighboring M12, where its expression is repressed by tey (PubMed:20504957). May also function in embryonic neuronal survival and the synaptic targeting of SNa motoneurons (PubMed:19018662).
(UniProt, P08953)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
Tl: Toll
Maternal expression of the Toll gene is required for the normal production and distribution of positional information in the embryo (Anderson et al., 1985); zygotic expression is required to maintain viability in early larvae (Gerttula et al., 1988). Toll mutants and deficiencies occurring in the mother result in lethal abnormalities in the pattern of gastrulation and the differentiation of cuticular structures in the offspring. When null alleles and deficiencies are homozygous in the zygote, delayed development and early lethality result. Females heterozygous for dominant Toll alleles are sterile, their lethal embryos being partially ventralized regardless of their genotype. Dorsoventral polarity is present; a furrow is formed in the midventral region, but the lateral cephalic fold is shifted to the dorsal side and the normal dorsal folds are missing. The cuticle lacks dorsal hairs, filzkorper, spiracles, head sensory organs, and a head skeleton; there are patches of denticles extending around the entire dorsoventral circumference of the embryo (Anderson et al., 1985a). The ventral nervous system is also expanded (Campos-Ortega, 1983). Embryos produced by females hemizygous for some dominant alleles (Tl1/Df; Tl3/Df) are ventralized, but the embryos of other hemizygotes (Tl2/Df; Tl4/Df) are dorsalized, all cells behaving at gastrulation and in differentiation like wild-type dorsal cells. In embryos derived from Tl/+ females, virtually the entire ectoderm capable of neurogenesis in response to absence of Dl function (Campos-Ortega, 1983, Wilhelm Roux's Arch. Dev. Biol. 192: 317-26). Whereas females heterozygous for recessive alleles of Tl are fertile, homozygous Tl-recessive females are viable but sterile, their lethal embryos lacking dorsoventral polarity and forming no ventral furrow at gastrulation. In most recessive alleles (Tlr5, Tlr6, Tlr7), the embryos are partially dorsalized with laterally derived structures (Anderson et al., 1985a); for example, Tlr6 embryos differentiate dorsal hairs, filzkorper, and ventral denticle bands of nearly normal width, but lack mesoderm (Anderson and Nusslein-Volhard, 1986). In one allele (Tlr4), however, embryos have no dorsal hairs and show rings of denticles as in TlD embryos (Anderson et al., 1985a). Hemizygotes for the Toll-recessives resemble the corresponding homozygotes in phenotype. A number of Toll alleles were obtained as reversions of the Toll-dominant phenotype (see table). When crossed to wildtype males, females heterozygous for a null-type reversion are fully fertile; however, when crossed to males who are also heterozygous for a Toll null, these females produce Tl-homozygotes who are zygotic lethals, dying as early larvae and producing no Toll transcript. Heteroallelic combinations of reversions such as Tlrv1/Tlrv2 produce sterile females with lethal dorsalized embryos. Females carrying combinations of certain reversions and Toll-dominant (or Toll-recessive) alleles produce embryos with phenotypes like those of Toll-dominant (or Toll-recessive) hemizygotes. Most of the reversions, when in trans to deficiencies, result in females with dorsalized embryos, but a few hemizygous reversion females (Tlrv21, Tlrv22, Tlrv23) produce ventralized embryos (Hashimoto et al., 1988). The lethal embryos of Df(3R)Tl-X/Df(3R)ro-XB3 (null) females (Hashimoto et al., 1988), are completely dorsalized, never making ventral furrows, filzkorper, or denticles; their germ bands fail to extend; no Toll transcript is produced in these embryos except when contributed by wild-type fathers (Gerttula et al., 1988). The 97D1-2 breakpoint of the Toll deficiency Df(3R)Tl-X maps within the 6.0 kb EcoRI fragment of a Toll clone (Hashimoto et al., 1988). Injection of wild-type cytoplasm into embryos of Toll-deficient females restores the wild-type dorsoventral pattern, the site of the injection determining the midventral part of the pattern (Anderson et al., 1985b); (also see molecular biology section).
Summary (Interactive Fly)
transmembrane receptor - IL1 homolog - a crucial protein for embryonic dorsal/ventral polarity and immunity - receptor for spätzle - activates the Tl intracellular signaling pathway
Gene Model and Products
Number of Transcripts
3
Number of Unique Polypeptides
2

Please see the GBrowse view of Dmel\Tl or the JBrowse view of Dmel\Tl 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 5.47
gene_with_stop_codon_read_through ; SO:0000697
Stop-codon suppression (UAG) postulated; FBrf0216884.
Gene model reviewed during 5.44
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)
FBtr0085059
5129
1097
FBtr0330154
5129
1117
FBtr0330155
4710
1097
Additional Transcript Data and Comments
Reported size (kB)
5.3 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0084431
124.7
1097
7.00
FBpp0303187
127.1
1117
6.78
FBpp0303188
124.7
1097
7.00
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

1097 aa isoforms: Tl-PB, Tl-PD
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)
In the absence of ligand, forms a low-affinity disulfide-linked homodimer (PubMed:24733933). In the presence of ligand, crystal structures show one Tl molecule bound to a spaetzle C-106 homodimer (PubMed:24282309, PubMed:24733933). However, the active complex probably consists of two Tl molecules bound to a spaetzle C-106 homodimer (PubMed:24282309, PubMed:24733933). This is supported by in vitro experiments which also show binding of the spaetzle C-106 dimer to 2 Tl receptors (PubMed:12872120). Ligand binding induces conformational changes in the extracellular domain of Tl (PubMed:24282309). This may enable a secondary homodimerization interface at the C-terminus of the Tl extracellular domain (PubMed:24282309).
(UniProt, P08953)
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\Tl using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (32 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Biological Process (20 terms)
Terms Based on Experimental Evidence (20 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from direct assay
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:Tl; FB:FBgn0262473
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from direct assay
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:Akt1; FB:FBgn0010379
inferred from genetic interaction with FLYBASE:Pi3K92E; FB:FBgn0015279
inferred from genetic interaction with FLYBASE:Pdk1; FB:FBgn0020386
inferred from genetic interaction with FLYBASE:InR; FB:FBgn0283499
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from direct assay
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN001555455
(assigned by GO_Central )
Cellular Component (7 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
inferred from high throughput direct assay
inferred from direct assay
(assigned by UniProt )
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN001555455
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN001555455
(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
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Expression pattern inferred from unspecified enhancer trap line.
Toll is expressed at high levels in embryos and pupae. Tl expression is noticably (2- to 5-fold) up-regulated in immune-challenged larvae and adults.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Protein, which is maternally provide is observed in the area between the somatic bud on the plasma membrane prior to embryonic cycle 13. At cellularization during cycle 14 the protein becomes concentrated at the basal membrane.
Tl protein is absent in the earliest stages of embryonic development, begins to accumulate prior to nuclear migration and peaks in late syncytial blastoderm embryos.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
inferred from high throughput direct assay
inferred from direct assay
(assigned by UniProt )
Expression Deduced from Reporters
Reporter: P{4xCME-lacZ}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{FZ}TlF336
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lwB}TlAK80
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Tl 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
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Images
Alleles, Insertions, and Transgenic Constructs
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
abdominal posterior fascicle & growth cone
filopodium & abdominal ventral longitudinal muscle 3
RP3 neuron & growth cone
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (16)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
2 of 15
No
Yes
 
2 of 15
No
Yes
 
2 of 15
No
Yes
 
2 of 15
No
Yes
2 of 15
No
Yes
2 of 15
Yes
Yes
2 of 15
Yes
Yes
2 of 15
Yes
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
Yes
No
 
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
 
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (14)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
5 of 15
Yes
Yes
3 of 15
No
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
Yes
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
Rattus norvegicus (Norway rat) (12)
3 of 13
Yes
Yes
2 of 13
No
No
2 of 13
No
No
2 of 13
No
Yes
2 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
No
1 of 13
No
No
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (11)
2 of 12
Yes
No
2 of 12
Yes
No
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
Danio rerio (Zebrafish) (21)
5 of 15
Yes
Yes
3 of 15
No
Yes
3 of 15
No
No
3 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
Yes
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
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
No
Caenorhabditis elegans (Nematode, roundworm) (2)
2 of 15
Yes
No
1 of 15
No
Yes
Arabidopsis thaliana (thale-cress) (7)
1 of 9
Yes
No
1 of 9
Yes
Yes
1 of 9
Yes
Yes
1 of 9
Yes
No
1 of 9
Yes
Yes
1 of 9
Yes
No
1 of 9
Yes
Yes
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) ( EOG091901OE )
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 persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091500U2 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Mayetiola destructor
Hessian fly
Mayetiola destructor
Hessian fly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Anopheles gambiae
Malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W00NQ )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Megachile rotundata
Alfalfa leafcutting bee
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Tribolium castaneum
Red flour beetle
Tribolium castaneum
Red flour beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X00MF )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G00ZN )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Ciona intestinalis
Vase tunicate
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (8)
4 of 10
3 of 10
3 of 10
2 of 10
2 of 10
2 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 ( 3 )
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 3 )
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?
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
RNA-protein
Physical Interaction
Assay
References
RNA-RNA
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
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
suppressible
External Data
Subunit Structure (UniProtKB)
In the absence of ligand, forms a low-affinity disulfide-linked homodimer (PubMed:24733933). In the presence of ligand, crystal structures show one Tl molecule bound to a spaetzle C-106 homodimer (PubMed:24282309, PubMed:24733933). However, the active complex probably consists of two Tl molecules bound to a spaetzle C-106 homodimer (PubMed:24282309, PubMed:24733933). This is supported by in vitro experiments which also show binding of the spaetzle C-106 dimer to 2 Tl receptors (PubMed:12872120). Ligand binding induces conformational changes in the extracellular domain of Tl (PubMed:24282309). This may enable a secondary homodimerization interface at the C-terminus of the Tl extracellular domain (PubMed:24282309).
(UniProt, P08953 )
Linkouts
DroID - A comprehensive database of gene and protein interactions.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Gene Group - Pathway Membership (FlyBase)
Toll-NF-KappaB Signaling Pathway Core Components -
In Drosophila, the canonical Toll signaling pathway is initiated by the binding of a spatzle ligand to Toll (Tl) or a Toll-like receptor leading to the nuclear localization of the NF-κB (dl or Dif) transcription factor. (Adapted from FBrf0091014 and FBrf0223077).
External Data
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3R
Recombination map
3-92
Cytogenetic map
Sequence location
3R:26,799,041..26,842,403 [+]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
97D2-97D2
Limits computationally determined from genome sequence between P{lacW}scribj7B3 and P{lacW}His2AvL1602
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
97D1-97D2
(determined by in situ hybridisation)
Location based on the breakpoints of several Tl revertant alleles.
97D-97D
(determined by in situ hybridisation)
Experimentally Determined Recombination Data
Left of (cM)
Right of (cM)
Notes
Tl7 and Tl8 map between e and ca, Tl9 maps between sr and e and Tl10 maps between sr and ca.
Stocks and Reagents
Stocks (25)
Genomic Clones (36)
cDNA Clones (101)
 

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
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.

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 database merge of
Source for merge of: Tl EP1051
Additional comments
A dorsalising activity for the heterologous ea, spz and Tl proteins in UV-ventralised Xenopus embryos is demonstrated: Tl dorsalises UV-treated X.laevis embryos. The activity is inhibited by co-injection of a dominant cact variant.
Other Comments
Embryonic extracts derived from a cross between Tl mutant mothers and Oregon R males have increased amounts of the 80 kD ea-Spn27A protein complex compared to wild type embryonic extracts.
dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.
RNAi screen using dsRNA made from templates generated with primers directed against this gene causes a phenotype when assayed in Kc167 and S2R+ cells: binucleate cells.
Tl requires only an endogenous protein ligand - the spz gene product - for activation and signalling. The mature, processed, form of the spz gene product binds to the Tl ectodomain with high affinity and with a stoichiometry of one spz dimer to two Tl receptors.
Tl regulates Drs expression through Dif.
nec negatively regulates the Tl signalling pathway.
Tl does not function as a pattern recognition receptor in the Drosophila host defence.
Tl acts in a cell autonomous manner in the fat body.
A linear activation cascade spz-Tl-cact-dl/Dif leads to the induction of the Drs gene in larval fat body cells.
Expression of Tl in a subset of epidermal cells, including the epidermal muscle attachment cells, but not in the musculature is necessary for proper muscle development in the embryo.
Tl mRNA is translationally activated by regulated cytoplasmic polyadenylation.
Targeting of either tub or pll product to the plasma membrane by myristylation is sufficient to activate the signal transduction pathway that leads to translocation of the dl product. Activated Tl induces a localized recruitment of tub and pll proteins to the plasma membrane.
Tl pathway is required for the nuclear import of dl in the immune response, but not required for the nuclear import of Dif. Cytoplasmic retention of both dl and Dif depends on cact protein. The two signalling pathways that target cact for degradation must discriminate between cact-dl and cact-Dif complexes.
Results from the expression of a constitutively activated form of the Tl receptor suggest that Tl signalling components diffuse in the plasma membrane or syncytial cytoplasm of the early embryo.
A combination of genetic manipulation and single-cell visualisation demonstrates the timing and cell specificity of muscular Tl expression can affect synaptogenesis of RP3 and other motoneuron growth cones.
The embryonic regulatory pathway, comprising the gene products between spz and cact (Tl, tub and pll) but not the genes acting upstream or downstream (ea and dl), is involved in the induction of the Drs gene in adults. Mutations that affect the synthesis of antimicrobial peptides dramatically lower the resistance of flies to infection.
Tl and pll can functionally interact to enhance dl activity synergistically.
An activated processed form of spz can activate Tl when injected into the extracellular space of early embryos (FBrf0074384). cact is rapidly degraded in response to spz injection.
Tl is one of several genes required for proper motoneuron and muscle specification. Loss of one or both copies of Tl leads to widespread defects in motoneuron number and muscle patterning.
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.
dl is not involved in the formation of melanotic tumours of Tl mutations.
tub is capable of acting as both a chaperon or escort for dl as it moves to the nucleus and then as a transcriptional coactivator. The intracytoplasmic domain of Tl is sufficient for activating the signalling pathway that leads to dl-tub nuclear translocation in Schneider cells.
Studies on a truncated Tl receptor indicate that the Tl receptor extracellular domain regulates the intrinsic signaling activity of its cytoplasmic domain.
dl is an embryonic phosphoprotein and its phosphorylation state is regulated by an intracellular signaling pathway initiated by the transmembrane receptor Tl. Using a combined genetic and biochemical approach it is demonstrated that activation of Tl stimulates an increase in the extent of dl phosphorylation.
Dorsal-ventral patterning is regulated by a signalling pathway that includes Tl and transcription factors, dl, that interact with related enhancers, rho. The κ enhancer from mouse is capable of generating lateral stripes of Ecol\lacZ gene expression in transgenic embryos in a pattern similar to that directed by rho enhancer. Results suggest that enhancers can couple conserved signalling pathways to divergent gene functions, dorso-ventral patterning and mammalian haematopoiesis.
The spz product acts immediately upstream of Tl in dorso-ventral pattern formation in the embryo, and may encode the ligand that activates Tl. The secreted spz product must be activated by proteolytic cleavage, and localized proteolytic processing of the spz protein determines where the receptor, Tl, is active.
Comparisons of early development to that in other insects have revealed conservation of some aspects of development, as well as differences that may explain variations in early patterning events.
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.
Sequence analysis of sim Tl and sli revealed a conserved sequence ACGTG that resembles the mammalian xenobiotic response element. This motif forms the core of an element required for CNS midline transcription.
Increased cytoplasmic calcium concentration and the expression of constitutively active Tl receptors can induce the relocalisation of dl in culture cells. Activation of endogenous Pka-C1, expression of wild type Tl receptors or treatment of cells with activators of Pkc53E and radical oxygen intermediates have only a marginal effects on the cellular distribution of dl protein.
Cytoplasmic injection studies indicate that the spatial information for the embryonic dorsal-ventral axis is largely derived from spatial cues in the extraembryonic compartment (most likely generated during oogenesis), which restrict the release of the putative Toll ligand. This ligand appears to originate from a ventrally restricted zone extending along the anterior-posterior axis, and its diffusion or graded release are required to determine the slope of the nuclear dorsal protein gradient. Both the Toll receptor and its ligand are in excess in wild type embryos.
Double mutant analysis indicates that ve acts upstream of Toll in dorsal-ventral axis formation, and the action of ve requires the grk-Egfr signaling pathway.
Analysis of Tl-Ecol\lacZ deletion constructs has identified a 750bp central nervous system midline enhancer in the Tl upstream region (between -21.kb and -1.4kb).
Double mutant combinations of Tl with ea alleles demonstrate that spatial regulation of ea activity by localized zymogen activation is a key initial event in defining the polarity of the dorsal-ventral embryonic pattern.
Toll enhances transport of dl protein into nucleus in cotransfected Schneider cells, perhaps via activated protein kinase A that phosphorylates dl gene product.
In addition to the Tl ligand, perivitelline fluid also contains three separate activities capable of rescuing ea, snk and spz. Serine proteolytic activity in the perivitelline fluid is required for the formation of the Tl ligand.
The cytoplasmic domain of the Tl protein is related to that of the human interleukin-1 receptor.
The properties of a peptide corresponding to residues 166-188 of the Tl protein have been studied in vitro.
The Tl protein is a glycoprotein which is tightly associated with embryonic membranes.
Recessive dorsalizing mutants of the dorsal group gene Tl have significantly reduced axial ratios in pupae.
Mutations in maternal dorsal class gene Tl do not interact with RpII140wimp.
Local activation of Tl by a Tl ligand initiates the formation of the dl nuclear concentration gradient, thereby determining the dorsoventral pattern.
The effects of an altered nucleocytoplasmic ratio on transcripts that normally undergo changes in transcript pattern in cell cycle 14 is studied. A delay in the maternal-to-zygotic transition of the dorsal-ventral polarity gene Tl is correlated with a decrease in nuclear density and a change in the cell cycle program.
sim gene product is required for the normal expression of Tl.
Involved in the regulatory hierarchy responsible for the asymmetric distribution and function of zygotic regulatory gene products along the DV axis of early embryos. Dominant cact mutants have a similar cuticle phenotype to that of zen- embryos.
Genetic and molecular analysis demonstrates that Tl is expressed and is functional zygotically as well as maternally.
Epistatic relationships exist between dorsalizing maternal effect mutations and "dppHin" alleles.
The expression of genes controlling neurogenesis is dependent on the previous activity of the genes controlling the development of the embryonic dorsal-ventral pattern. Double mutants N55e11 and Dl9P with Tl had neuralization of the entire ectoderm, a huge CNS and no epidermis as it had been substituted for by neural tissue.
Females carrying the dominant allele Tl3, when combined with the mutants gd, ndl, pip, snk, or ea, produce embryos that are lateralized like embryos derived from Tlrv8 females; these embryos lack dorsalmost and ventralmost pattern elements and have rings of denticles (Anderson, Jurgens and Nusslein-Volhard, 1985). Some alleles of ea increase the probability that the temperature-sensitive alleles Tlr5, Tlr6 and Tlr7 will survive. An interaction has been reported between the recessive allele Tlr7 and dpp (Irish and Gelbart, 1987). Double mutants of Tl3 and dl produce embryos that are completely dorsalized and indistinguishable from the embryos of dl homozygotes. Females carrying Tl2 or Tl4 in combination with gd, ndl, or dl also produce dorsalized embryos.
Maternal expression of the Toll gene is required for the normal production and distribution of positional information in the embryo (Anderson, Jurgens and Nusslein-Volhard, 1985; Anderson, Bokla and Nusslein-Volhard, 1985); zygotic expression is required to maintain viability in early larvae (Gerttula, Jin and Anderson, 1988). Toll mutants and deficiencies occurring in the mother result in lethal abnormalities in the pattern of gastrulation and the differentiation of cuticular structures in the offspring. When null alleles and deficiencies are homozygous in the zygote, delayed development and early lethality result. Females heterozygous for dominant Toll alleles are sterile, their lethal embryos being partially ventralized regardless of their genotype. Dorsoventral polarity is present; a furrow is formed in the midventral region, but the lateral cephalic fold is shifted to the dorsal side and the normal dorsal folds are missing. The cuticle lacks dorsal hairs, filzkorper, spiracles, head sensory organs and a head skeleton; there are patches of denticles extending around the entire dorsoventral circumference of the embryo (Anderson, Jurgens and Nusslein-Volhard, 1985). The ventral nervous system is also expanded (Campos-Ortega, 1983). Embryos produced by females hemizygous for some dominant alleles (Tl1/Df; Tl3/Df) are ventralized, but the embryos of other hemizygotes (Tl2/Df; Tl4/Df) are dorsalized, all cells behaving at gastrulation and in differentiation like wild-type dorsal cells. In embryos derived from Tl/+ females, virtually the entire ectoderm capable of neurogenesis in response to absence of Dl function (Campos-Ortega, 1983). Whereas females heterozygous for recessive alleles of Tl are fertile, homozygous Tl-recessive females are viable but sterile, their lethal embryos lacking dorsoventral polarity and forming no ventral furrow at gastrulation. In most recessive alleles (Tlr5, Tlr6, Tlr7), the embryos are partially dorsalized with laterally derived structures (Anderson, Jurgens and Nusslein-Volhard, 1985); for example, Tlr6 embryos differentiate dorsal hairs, filzkorper and ventral denticle bands of nearly normal width, but lack mesoderm (Anderson and Nusslein-Volhard, 1986). In one allele (Tlr4), however, embryos have no dorsal hairs and show rings of denticles as in TlD embryos (Anderson, Jurgens and Nusslein-Volhard, 1985). Hemizygotes for the Toll-recessives resemble the corresponding homozygotes in phenotype. A number of Toll alleles were obtained as reversions of the Toll-dominant phenotype. When crossed to wild-type males, females heterozygous for a null-type reversion are fully fertile; however, when crossed to males who are also heterozygous for a Toll null, these females produce Tl-homozygotes who are zygotic lethals, dying as early larvae and producing no Toll transcript. Heteroallelic combinations of reversions such as Tlrv1/Tlrv2 produce sterile females with lethal dorsalized embryos. Females carrying combinations of certain reversions and Toll-dominant (or Toll-recessive) alleles produce embryos with phenotypes like those of Toll-dominant (or Toll-recessive) hemizygotes. Most of the reversions, when in trans to deficiencies, result in females with dorsalized embryos, but a few hemizygous reversion females (Tlrv21, Tlrv22, Tlrv23) produce ventralized embryos (Hashimoto et al., 1988). The lethal embryos of Df(3R)Tl-X/Df(3R)ro-XB3 (null) females (Hashimoto, Hudson and Anderson, 1988), are completely dorsalized, never making ventral furrows, filzkorper, or denticles; their germ bands fail to extend; no Toll transcript is produced in these embryos except when contributed by wild-type fathers (Gerttula, Jin and Anderson, 1988). The 97D1-2 breakpoint of the Toll deficiency Df(3R)Tl-X maps within the 6.0 kb EcoRI fragment of a Toll clone (Hashimoto, Hudson and Anderson, 1988). Injection of wild-type cytoplasm into embryos of Toll-deficient females restores the wild-type dorsoventral pattern, the site of the injection determining the midventral part of the pattern (Anderson, Bokla and Nusslein-Volhard, 1985).
Origin and Etymology
Discoverer
Wieschaus and Nusslein-Volhard.
Etymology
Identification
External Crossreferences and Linkouts ( 74 )
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.
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
UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
Other crossreferences
Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
FlyMine - An integrated database for Drosophila genomics
GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
KEGG Genes - Molecular building blocks of life in the genomic space.
KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
modMine - A data warehouse for the modENCODE project
Synonyms and Secondary IDs (25)
Reported As
Symbol Synonym
Fs(3)Tl
Tl
(Araki et al., 2019, Chowdhury et al., 2019, Pippi et al., 2019, Wurster et al., 2019, Xu et al., 2019, Alpar et al., 2018, Green et al., 2018, Merkel et al., 2018, Segal and Frenkel, 2018, Christesen et al., 2017, Duneau et al., 2017, Early et al., 2017, Koenecke et al., 2017, Louradour et al., 2017, Park et al., 2017, Rohde et al., 2017, Lim et al., 2016, Liu et al., 2016, Schmid et al., 2016, Aradska et al., 2015, Yamamoto-Hino et al., 2015, Zang et al., 2015, Zhan et al., 2015, Arvanitis et al., 2014, Farmakiotis et al., 2014, Ferreira et al., 2014, Ocorr et al., 2014, Taylor et al., 2014, Tevy et al., 2014, Zhou et al., 2014, Ben-Ami et al., 2013, McIlroy et al., 2013, Ozkan et al., 2013, Ozkan et al., 2013, Ozkan et al., 2013, Samaraweera et al., 2013, Webber et al., 2013, Zanette et al., 2013, Broderick et al., 2012, Bryantsev and Cripps, 2012, Cho et al., 2012, Haskel-Ittah et al., 2012, Lemaitre et al., 2012, Jungreis et al., 2011, Marcu et al., 2011, Venken et al., 2011, Wang et al., 2011, Anbutsu and Fukatsu, 2010, Chen et al., 2010, Goto et al., 2010, Hill-Burns and Clark, 2010, Inaki et al., 2010, Kim et al., 2010, Kong et al., 2010, Lund et al., 2010, Stein et al., 2010, Valanne et al., 2010, Yagi et al., 2010, Ahmad et al., 2009, Buchon et al., 2009, Buchon et al., 2009, Chamilos et al., 2009, Hashimoto et al., 2009, Jin et al., 2009, Lamaris et al., 2009, Obbard et al., 2009, Roeder et al., 2009, Tan et al., 2009, Chamilos et al., 2008, Davis et al., 2008, Gilchrist et al., 2008, Markstein et al., 2008, Tan et al., 2008, Vonkavaara et al., 2008, Witzberger et al., 2008, Zhu et al., 2008, Minidorff et al., 2007, Morozova et al., 2007, Nehme et al., 2007, Shen and Tanda, 2007, Taylor and Kimbrell, 2007, Wu et al., 2007, Xing et al., 2007, Zeitlinger et al., 2007, Zeitouni et al., 2007, Zhang et al., 2007, Brun et al., 2006, Carneiro et al., 2006, Kambris et al., 2006, Keranen et al., 2006, Kim et al., 2006, LeMosy, 2006, Mizutani et al., 2006, Senger et al., 2006, Castillejo-Lopez and Häcker, 2005, Mizutani et al., 2005, Thoetkiattikul et al., 2005, Wang et al., 2005, Wertheim et al., 2005, Alarco et al., 2004, Cowden and Levine, 2003, Lau et al., 2003, Leulier et al., 2003, Rutschmann et al., 2002, Tauszig-Delamasure et al., 2002, Araujo and Bier, 2000)
mat(3)9
mel(3)10
mel(3)9
Name Synonyms
EP1051
Protein toll precursor
Toll
(Mehmeti et al., 2019, Meltzer, 2019-, Meltzer et al., 2019, Valanne et al., 2019, Abed et al., 2018, Coll et al., 2018, Katsukawa et al., 2018, Merkel et al., 2018, Millet-Boureima et al., 2018, Min and Tatar, 2018, Palmer et al., 2018, Parvy et al., 2018, Richardson and Portela, 2018, Schmidt and Grosshans, 2018, Segal and Frenkel, 2018, Sherri et al., 2018, Yu et al., 2018, Del Signore et al., 2017, Hao and Jin, 2017, Kashio et al., 2017, Morales and Li, 2017, Mussabekova et al., 2017, Trinder et al., 2017, Eichhorn et al., 2016, Faye and Lindberg, 2016, Mistry et al., 2016, Rahimi et al., 2016, Vanha-Aho et al., 2016, Yamamoto-Hino and Goto, 2016, Kanoh et al., 2015, Liu et al., 2015, Lucas et al., 2015, Shaukat et al., 2015, Wu et al., 2015, Yamamoto-Hino et al., 2015, Bonnay et al., 2014, Carvalho et al., 2014, Farmakiotis et al., 2014, Ferreira et al., 2014, Hultmark and Szabad, 2014.5.19, Imler, 2014, Ji et al., 2014, Keebaugh and Schlenke, 2014, Kurata, 2014, Lee and Hyun, 2014, Lindsay and Wasserman, 2014, Myllymäki et al., 2014, Ocorr et al., 2014, Panayidou et al., 2014, Parthier et al., 2014, Salazar-Jaramillo et al., 2014, Ben-Ami et al., 2013, Cui et al., 2013, Ferrandon, 2013, Gueguen et al., 2013, Kingsolver et al., 2013, Lagha et al., 2013, Mbodj et al., 2013, Nelson et al., 2013, Rancès et al., 2013, Samaraweera et al., 2013, Stelter et al., 2013, Zanette and Kontoyiannis, 2013, Zanette and Kontoyiannis, 2013, Zanette et al., 2013, Bryantsev and Cripps, 2012, Chopra et al., 2012, Haskel-Ittah et al., 2012, Lemaitre et al., 2012, Nakamoto et al., 2012, Vanha-Aho et al., 2012, Chtarbanova and Imler, 2011, Clark et al., 2011, Dunipace et al., 2011, Fulkerson and Estes, 2011, Galac and Lazzaro, 2011, Garcia and Stathopoulos, 2011, Lynch and Roth, 2011, Marcu et al., 2011, Nehme et al., 2011, Valanne et al., 2011, Vazquez-Pianzola et al., 2011, Wang et al., 2011, Yano and Kurata, 2011, Arnot et al., 2010, Ben-Ami et al., 2010, Chamilos et al., 2010, Chen et al., 2010, Coll et al., 2010, Huang et al., 2010, Inaki et al., 2010, Junell et al., 2010, Kim et al., 2010, Kurata, 2010, Kuttenkeuler et al., 2010, Lund et al., 2010, Sabin et al., 2010, Stein et al., 2010, Tanji et al., 2010, Valanne et al., 2010, Yagi et al., 2010, Ahmad et al., 2009, Buchon et al., 2009, Buchon et al., 2009, Chamilos et al., 2009, Cronin et al., 2009, Diangelo et al., 2009, Jin et al., 2009, Liu et al., 2009, Mavrakis et al., 2009, Zsindely et al., 2009, Araujo et al., 2008, Baudot et al., 2008, Cao et al., 2008, Cui et al., 2008, DiAngelo et al., 2008, Gordon et al., 2008, Goto et al., 2008, Hong et al., 2008, Kleino et al., 2008, Qi et al., 2008, Stein et al., 2008, Stramer et al., 2008, Tanda et al., 2008, Tan et al., 2008, Tsai et al., 2008, Vonkavaara et al., 2008, Witzberger et al., 2008, Wu and Sato, 2008, Zhu et al., 2008, Busse et al., 2007, Fenckova and Dolezal, 2007, Inaki et al., 2007, Kuranaga and Miura, 2007, Kuttenkeuler and Boutros, 2007, Morozova et al., 2007, Nehme et al., 2007, Pal et al., 2007, Pham et al., 2007, Scherfer et al., 2007, Tanji et al., 2007, Tao et al., 2007, Tao et al., 2007, Tao et al., 2007, Taylor and Kimbrell, 2007, Waterhouse et al., 2007, Williams et al., 2007, Wu et al., 2007, Xing et al., 2007, Zeitlinger et al., 2007, Zeitlinger et al., 2007, Zeitlinger et al., 2007, Zeitouni et al., 2007, Akira et al., 2006, Araujo et al., 2006, Biemar et al., 2006, Brun et al., 2006, Chen, 2006, Chen et al., 2006, Davidson and Erwin, 2006, Guan et al., 2006, Jang et al., 2006, Kambris et al., 2006, Keranen et al., 2006, LeMosy, 2006, Minakhina and Steward, 2006, Mulinari et al., 2006, Sambandan et al., 2006, Scherfer et al., 2006, Shi et al., 2006, Ulvila et al., 2006, Zaffran et al., 2006, Zimmermann et al., 2006, Burnette et al., 2005, Gesellchen et al., 2005, Mace et al., 2005, Pal and Wu, 2005, Staudt et al., 2005, Wang et al., 2005, Yagi and Ip, 2005, Zhou et al., 2005, Roxstrom-Lindquist et al., 2004, Cowden and Levine, 2003, Leulier et al., 2003, Park et al., 2003, Ritzenthaler and Chiba, 2003, Christophides et al., 2002, Ooi et al., 2002, Rutschmann et al., 2002, Tauszig-Delamasure et al., 2002, Furlong et al., 2001, Horng and Medzhitov, 2001, Luo et al., 2001, Araujo and Bier, 2000, Rutschmann et al., 2000, Meng et al., 1999, Braun et al., 1998, Govind et al., 1998, Uttenweiler-Joseph et al., 1998, Ferrandon et al., 1997, Ntwasa et al., 1997, Schisa and Strickland, 1996, Rosetto et al., 1995, Wharton and Crews, 1993, Leptin et al., 1992, Schupbach and Wieschaus, 1986)
Secondary FlyBase IDs
  • FBgn0003717
  • FBgn0062707
Datasets (0)
Study focus (0)
Experimental Role
Project
Project Type
Title
References (898)