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Gene Dmel\Tl

General Information
Symbol	Dmel\Tl	Species	D. melanogaster
Name	Toll	Annotation symbol	CG5490
Feature type	protein_coding_gene	FlyBase ID	FBgn0262473
Gene Model Status	Current	Stock availability	22 publicly available
Also Known As	Toll, Toll-1, T1, dToll
Genomic Location
Chromosome (arm)	3R	Recombination map	3-91
Cytogenetic map	97D2-97D2	Sequence location	3R:22,624,765..22,668,125 [+]
Genomic Maps Select View:Gene Models/EvidenceExpression/RegulationMegaViewGene Disruptions, Stocks and Reagents modENCODE GBrowse	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene Toll is referred to in FlyBase by the symbol Dmel\Tl (CG5490, FBgn0262473). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: protein binding; TIR domain binding. There is experimental evidence for 12 unique biological process terms, many of which group under: immune system process; immune response; biological regulation; system development; positive regulation of cellular biosynthetic process; cellular process; positive regulation of cellular metabolic process; cellular component organization or biogenesis; cardiovascular system development; neuron differentiation; dorsal/ventral axis specification. 169 alleles are reported. The phenotypes of these alleles are annotated with: organ system; multicellular structure; embryonic/larval hemocoel; portion of tissue; organ system subdivision; anatomical structure; extended germ band embryo; non-connected developing system; peripheral nervous system; embryonic dorsal vessel; embryonic/larval neuron; spiracle; late extended germ band embryo. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: Cysteine-rich flanking region, C-terminal; Interleukin-1 receptor, type I/Toll precursor; Leucine-rich repeat; Leucine-rich repeat, typical subtype; Leucine-rich repeat-containing N-terminal; Toll/interleukin-1 receptor homology (TIR) domain. Gene sequence location is 3R:22624765..22668125.
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	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).
Recent Updates
Description	What does this section display? This section contains items that were added to this record for each release. It currently only tracks new links between this FlyBase report and other FlyBase data classes (e.g. genes, references, stocks) or controlled vocabulary terms (e.g. GO, anatomy terms). What does this section not display? This section does not currently display links that were removed or gene model changes.
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FB2012_01	References Kim et al., 2010, Exp. Gerontol. 45(12): 984--987 Controlled Vocabulary Terms heart development Sequence features BKN45150 HFA16885 BKN23697
FB2011_10	References Clark et al., 2011, Curr. Biol. 21(19): 1672--1677 Vazquez-Pianzola et al., 2011, Dev. Biol. 357(2): 404--418
All updates	Click here to see a list of all updates to this record from FB2010_08 and on.
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 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   Location based on the breakpoints of several Tl revertant alleles.   (Tearle and Nusslein-Volhard, 1987) 97D1-97D2   (determined by in situ hybridisation)   (Ranz et al., 2001) 97D-97D   (determined by in situ hybridisation)   (Nose et al., 1992)
Experimentally Determined Recombination Data
Location	3-91 (Tearle and Nusslein-Volhard, 1987)
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. (Erdelyi and Szabad, 1989)
Gene Model & Products
Please see the GBrowse view of Dmel\Tl for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
	DGC clone appears problematic (GH03720): incomplete CDS; DGC:RE46574 okay.
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) Tl-RA FBtr0085060  NM_170287   5517   1097 Tl-RB FBtr0085059  NM_079794   5127   1097
Additional Transcript Data & Comments
Reported size (kB)	5.3 (northern blot) (Tauszig et al., 2000)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein Tl-PA   FBpp0084432   124.7   1097   7.00     NP_733166   AAF56624 Tl-PB   FBpp0084431   124.7   1097   7.00     NP_524518   AAN14086
Additional Polypeptide Data & Comments
Reported size (kDa)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Toll/interleukin-1 receptor homology (TIR) domain (IPR000157) Leucine-rich repeat-containing N-terminal (IPR000372) Cysteine-rich flanking region, C-terminal (IPR000483) Leucine-rich repeat (IPR001611) Leucine-rich repeat, typical subtype (IPR003591) Interleukin-1 receptor, type I/Toll precursor (IPR004075)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA202072   AA202511   AA263933   AA541030   AE014297 AAF56624   AE014297 AAN14086   AI063746   AI519197   AQ025161   AQ026361   AW940139   AY121616 AAM51943   AY349649 AAQ64932   AY349650 AAQ64933   AY349651 AAQ64934   AY349652 AAQ64935   AY349653 AAQ64936   AY349654 AAQ64937   AY349655 AAQ64938   BI164942   BI168001   BI171729   BI227539   BI229395   BI241665   BI361602   BI362102   BI362120   BI577715   BI579422   BT011023   BT031153   BT125764   BT126067   CO177276   CO178876   CO190636   CO263943   CO269176   CO270524   CO275104   CO277963   CO281524   CO281857   CO283182   CO284788   CO289626   CO290014   CO294601   CO296587   CO304238   CO311790   CO312992   CO313422   CO314284   CO315158   CO317052   CO317375   CO320667   CO326611   CO331439   CZ480900   DN154477   EC058030   EC058188   EC058797   EC063895   EC065129   EC065244   EC065386   EC072230   EC076938   EC081183   EC089483   EC089523   EC090070   EC090487   EC207754   EC209936   EC209944   EC211316   EC216394   EC219759   EC221187   EC222334   EC224792   EC228557   EC230100   EC233282   EC234280   EC244011   EC245676   EC247657   EC247904   EC256001   EC256786   EC258751   EV575273   GH942726   GH943655   GH947941   GH951590   GH956334   GH956885   GH958625   GH961180   GH961304   GH961317   GH969442   GH970791   GH972698   GH979542   GH979597   GH980609   GH982485   M19969 AAA28941
UniProtKB/Swiss-Prot	P08953
UniProtKB/TrEMBL
Mapped Features
Mapped Features have been reorganized, please see this article for details. Additional mapped features and mutations can be found on GBrowse or related reports. Type Symbol & Location Additional Notes References
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage 14 -- 16   embryonic cardioblast   (Wang et al., 2005) embryonic stage 16   embryonic dorsal vessel   (Wang et al., 2005) embryonic/larval aorta   (Wang et al., 2005) heart primordium   (Wang et al., 2005) northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage & pupal stage     (Tauszig et al., 2000)
Additional Descriptive Data
	Dvir\Tl 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. (Tauszig et al., 2000)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
immunolocalization Stage Tissue/Position (including subcellular localization) Reference embryonic cycle 1 -- embryonic cycle 12     • plasma membrane (Lecuit et al., 2002) embryonic stage 4     (Gay and Keith, 1992) embryonic cycle 14     • plasma membrane | basal (Lecuit et al., 2002) embryonic stage 14 -- 16   embryonic cardioblast   (Wang et al., 2005) embryonic stage 16   embryonic dorsal vessel   (Wang et al., 2005) embryonic/larval aorta   (Wang et al., 2005) heart primordium   (Wang et al., 2005)
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. (Gay and Keith, 1992, Lecuit et al., 2002) 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. (Gay and Keith, 1992)
Marker for
Subcellular Localization
CV Term	plasma membrane (Gay and Keith, 1992)
Notes
High-Throughput Expression Data
or
See Gelbart and Emmert, 2010.10.13 for analysis details and data files for all genes. modENCODE Temporal Expression Data (Graveley et al., 2011) FlyAtlas Anatomical Expression Data (Chintapalli et al., 2007)
Expression Clusters
	A cluster of genes with similar mRNA expression dynamics across development. (The modENCODE Consortium, 2010) mE2_34_mRNA_expression_cluster_17
External Data & Images
Linkouts
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal | dominant | embryonic stage Tl4 lethal | dominant | embryonic stage | maternal effect Tlr1 lethal | dominant | maternal effect Tl1 Tl3 Tl4 lethal | embryonic stage | germline clone | non-rescuable maternal effect | recessive Tl3R-39-38 Tl3R-50-33 Tl3R-209-21 Tl3R-267-38 lethal | embryonic stage | maternal effect Tl6 Tl7 Tl8 Tl9 Tl10 Tlr2 Tlr3 lethal | embryonic stage | maternal effect | recessive Tlr3 Tlr4 TlrK344 lethal | maternal effect Tl8 Tlr5 lethal | maternal effect | recessive Tlr6 Tlr7 lethal | maternal effect | recessive | heat sensitive Tlr1 Tlr2 Tlr3 Tlr4 lethal | recessive Tl1 Tl2 Tl3 viable TlMhc.PR Sterility Allele female sterile | dominant Tl3 female sterile germline-dependent | dominant Tl3 Other Phenotypes Allele decreased cell death | embryonic stage 17, with Scer\GAL4elav-C155 Tl10b.Scer\UAS immune response defective Tl3 Tlr3 immune response defective (with Tlr3) Tlrv1 immune response defective (with Tlrv1) Tlr3 melanotic mass phenotype | dominant Tl8 neuroanatomy defective TlMhc.PR neuroanatomy defective | embryonic stage TlMhc.PR neuroanatomy defective | embryonic stage, with Scer\GAL4tey-5053A TlEP1051 stress response defective Tl8 visible | heat sensitive Tlr2 Tlr3 wild-type Tl1.4 Phenotype manifest in Allele abdominal posterior fascicle & growth cone TlMhc.PR abdominal segment motor neuron TlMhc.PR abdominal segment motor neuron, with Scer\GAL4tey-5053A TlEP1051 amnioserosa (with Tlr2) Tlrv19 amnioserosa (with Tlrv19) Tlr2 cardioblast (with Tlr3) Tlr4 cardioblast (with Tlr4) Tlr3 crystal cell, with Scer\GAL4He.PZ Tl10b.Scer\UAS denticle belt, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 Tl10b-Nar.Scer\UAS Tl10b.Scer\UAS Tl10b-Nae.Scer\UAS Tl10b-3559.Scer\UAS dorsal ectoderm (with Tlr2) Tlrv19 dorsal ectoderm (with Tlrv19) Tlr2 embryo Tl3 Tl8 embryo | dorsal ventral gradient (with Tlr3) TlrK344 embryo | dorsal ventral gradient (with TlrK344) Tlr3 embryonic/first instar larval cuticle | dorsal Tl3 embryonic/first instar larval cuticle | germline clone | maternal effect Tl3R-209-21 embryonic/first instar larval cuticle | maternal effect Tl3 embryonic/larval cuticle Tl8 embryonic/larval hemocyte Tl8 embryonic/larval hemocyte (with Tlr3) Tlrv1 embryonic/larval hemocyte (with Tlrv1) Tlr3 embryonic/larval hemocyte | supernumerary Tl8 embryonic/larval hemolymph Tl8 embryonic/larval lymph gland Tl8 embryonic dorsal epidermis Tl4 Tl1 Tl3 Tl2 embryonic dorsal epidermis | maternal effect Tl3 embryonic dorsal vessel (with Tlr3) Tlr4 embryonic dorsal vessel (with Tlr4) Tlr3 embryonic epidermis Tl3 embryonic head TlDB1 embryonic leading edge cell (with Tlr2) Tlrv19 embryonic leading edge cell (with Tlrv19) Tlr2 embryonic ventral epidermis Tl4 Tl1 Tl3 Tl2 embryonic ventral epidermis | maternal effect Tl3 extended germ band embryo TlDB1 filopodium & abdominal ventral longitudinal muscle 3 TlMhc.PR filzkorper, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 Tl10b-Nar.Scer\UAS Tl10b.Scer\UAS Tl10b-Nae.Scer\UAS Tl10b-3559.Scer\UAS hemocyte, with Scer\GAL4He.PZ Tl10b.Scer\UAS hemocyte | larval stage Tl8 lamellocyte Tl8 lamellocyte, with Scer\GAL4He.PZ Tl10b.Scer\UAS lamellocyte | ectopic Tl8 lamellocyte | supernumerary Tl8 lamellocyte | supernumerary, with Scer\GAL4Cg.PA Tl10b.Scer\UAS.T:Zzzz\FLAG melanotic mass Tl3 Tl8 melanotic mass, with Scer\GAL4He.PZ Tl10b.Scer\UAS mesoderm Tl8 mesoderm | maternal effect Tl8 plasmatocyte Tl8 plasmatocyte | supernumerary Tl8 plasmatocyte | supernumerary, with Scer\GAL4Cg.PA Tl10b.Scer\UAS.T:Zzzz\FLAG podocyte Tl8 RP1 neuron | heat sensitive Tlr2 Tlr1 RP3 neuron TlMhc.PR RP3 neuron & growth cone TlMhc.PR RP3 neuron | heat sensitive Tlr2 Tlr1 RP4 neuron | heat sensitive Tlr2 Tlr1 sessile hemocyte, with Scer\GAL4Hml.Δ Tl10b.Scer\UAS
Classical Alleles ( 65 )
For All Classical Alleles Show
Allele of Tl Class Mutagen Stocks Known lesion Tl3 gain of function allele ethyl methanesulfonate 2 Yes Tlr3 hypomorphic allele - genetic evidence ethyl methanesulfonate 2 -- Tlr4 loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate 2 Yes Tlrv18 loss of function allele, amorphic allele - genetic evidence X ray 1 Yes Tl8 gain of function allele ethyl methanesulfonate 1 Yes Tle03133 piggyBac transposase 1 -- TlKG03609 P-element activity 1 -- TlMI00181 1 -- TlMI01254 1 -- TlNP0756 1 -- TlrK344 P-element activity 1 -- Tl2 amorphic allele - genetic evidence, gain of function allele ethyl methanesulfonate 0 Yes TlreQ amorphic allele - genetic evidence 0 -- Tlrv13 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate 0 -- Tlrv16 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate 0 -- Tlrv19 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate 0 -- Tlrv1 loss of function allele, amorphic allele - genetic evidence X ray 0 -- Tlrv24 amorphic allele - genetic evidence 0 -- Tlrv25 amorphic allele - genetic evidence 0 -- Tlrv2 amorphic allele - genetic evidence X ray 0 -- Tlrv4 amorphic allele - genetic evidence, loss of function allele X ray 0 -- Tl10 ethyl methanesulfonate 0 -- Tl1 gain of function allele ethyl methanesulfonate 0 Yes Tl3p Delta2-3 0 Yes Tl3R-209-21 ethyl methanesulfonate 0 -- Tl3R-267-38 ethyl methanesulfonate 0 -- Tl3R-39-38 ethyl methanesulfonate 0 -- Tl3R-50-33 ethyl methanesulfonate 0 -- Tl3t Delta2-3 0 Yes Tl4 gain of function allele ethyl methanesulfonate 0 Yes Tl5 P-element activity 0 -- Tl6 gain of function allele ethyl methanesulfonate 0 Yes Tl7 ethyl methanesulfonate 0 -- Tl9 ethyl methanesulfonate 0 -- Tl9BREQ 0 -- Tl9QRE1 0 -- Tl9QURE 0 -- TlAK80 P-element activity 0 -- TlDB1 gain of function allele 0 Yes TlDB2 gain of function allele 0 Yes TlDB3 gain of function allele 0 Yes TlEP1051 P-element activity 0 -- TlF336 P-element activity 0 Yes Tlr1 loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes Tlr2 loss of function allele, hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes Tlr5 antimorphic allele - genetic evidence ethyl methanesulfonate 0 -- Tlr6 loss of function allele ethyl methanesulfonate 0 Yes Tlr7 loss of function allele ethyl methanesulfonate 0 Yes Tlrv10 loss of function allele ethyl methanesulfonate 0 -- Tlrv11 loss of function allele ethyl methanesulfonate 0 -- Tlrv12 loss of function allele ethyl methanesulfonate 0 -- Tlrv14 loss of function allele ethyl methanesulfonate 0 -- Tlrv15 loss of function allele ethyl methanesulfonate 0 -- Tlrv17 loss of function allele ethyl methanesulfonate 0 -- Tlrv20 X ray 0 -- Tlrv21 loss of function allele 0 -- Tlrv22 loss of function allele PM hybrid dysgenesis 0 Yes Tlrv23 loss of function allele 0 -- Tlrv3 X ray 0 -- Tlrv5 X ray 0 -- Tlrv6 X ray 0 -- Tlrv7 loss of function allele ethyl methanesulfonate 0 -- Tlrv8 loss of function allele ethyl methanesulfonate 0 -- Tlrv9 loss of function allele ethyl methanesulfonate 0 -- Tlunspecified 0 --
Alleles Carried on Transgenic Constructs ( 104 )
For All Alleles Carried on Transgenic Constructs Show
Allele of Tl Class Mutagen Stocks Known lesion TlScer\UAS.P\T.T.T:Avic\GFP-PA in vitro construct - coding region fusion in vitro construct - regulatory fusion 2 Yes TlScer\UAS.P\T.T:Avic\GFP-YFP.Venus in vitro construct - coding region fusion in vitro construct - regulatory fusion 2 Yes TlGL00474 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes TlJF01276 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes TlJF01491 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes TlKK103505 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes Tl1.4 in vitro construct - regulatory fusion 0 Yes Tl1.cSa in vitro construct 0 Yes Tl10b-3559.Scer\UAS in vitro construct - deletion 0 Yes Tl10b-Nae.Scer\UAS in vitro construct - deletion 0 Yes Tl10b-Nar.Scer\UAS in vitro construct - deletion 0 Yes Tl10b-Stu.Scer\UAS in vitro construct - deletion 0 Yes Tl10b-ΔPEST.Scer\UAS in vitro construct - deletion 0 Yes Tl10b.Act5C in vitro construct - amino acid replacement in vitro construct - regulatory fusion 0 Yes Tl10B.hs in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement in vitro construct - regulatory fusion 0 Yes Tl10B.MtnA.T:SV5\V5 in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - site directed mutagenesis 0 Yes Tl10b.Scer\UAS.cYa 0 Yes Tl10b.Scer\UAS.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - amino acid replacement 0 Yes Tl10b.Scer\UAS.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - amino acid replacement 0 Yes Tl10b.Scer\UAS in vitro construct - regulatory fusion 0 Yes Tl10b.ΔC1.Scer\UAS.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - amino acid replacement 0 Yes Tl10b.ΔC2.Scer\UAS.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - amino acid replacement 0 Yes Tl520.Act5C in vitro construct - deletion 0 Yes Tl6.5 in vitro construct - regulatory fusion 0 Yes Tl8.bcd.3'UTR.bcd in vitro construct - regulatory fusion 0 Yes Tl8.bcd.3'UTR.Hsp83 in vitro construct - regulatory fusion 0 Yes Tl8.cSa in vitro construct 0 Yes TlAct5C.PN in vitro construct - regulatory fusion 0 Yes TlBstXI.Act5C in vitro construct - deletion 0 Yes TlC755Y.C757Y.C781Y.C799Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC755Y.C757Y.C781Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC755Y.C757Y.C799Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC755Y.C757Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC755Y.C781Y.C799Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC755Y.C781Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC755Y.C799Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC755Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC757Y.C781Y.C799Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC757Y.C781Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC757Y.C799Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC757Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781D.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781E.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781G.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781H.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781L.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781P.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781R.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781S.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781stop.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781T.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781W.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781Y.C799Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC781Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlC799Y.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlDK.BB4 in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes TldsRNA.3289-3866 in vitro construct - RNAi 0 Yes TldsRNA.cWa in vitro construct - RNAi 0 Yes TldsRNA.cZa in vitro construct - RNAi 0 Yes Tlfl.cSa in vitro construct 0 Yes TlGD305 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes TlGV.BB8 in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes Tlhs.F9 in vitro construct - regulatory fusion 0 Yes TlIC-NaeI.Act5C.T:Ivir\HA1 in vitro construct - regulatory fusion 0 Yes TlIC-StuI.Act5C.T:Ivir\HA1 in vitro construct - regulatory fusion 0 Yes TlIC.Act5C.T:Ivir\HA1 in vitro construct - regulatory fusion 0 Yes TlMhc.PR in vitro construct - regulatory fusion 0 Yes TlNaeI.Act5C in vitro construct - deletion 0 Yes TlNarI.Act5C in vitro construct - deletion 0 Yes TlRV.Act5C in vitro construct - amino acid replacement 0 Yes TlScer\UAS.cHa in vitro construct - regulatory fusion 0 Yes TlScer\UAS.cMa in vitro construct - regulatory fusion 0 Yes TlScer\UAS.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes TlScer\UAS.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes TlStuI.Act5C in vitro construct - deletion 0 Yes Tltrunc1 in vitro construct 0 Yes TlVE.BB7 in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes TlVH.BB7 in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes TlVP.BB7 in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes TlVR.BB7 in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes TlΔIC in vitro construct 0 Yes TlΔLRR.MtnA in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes TlΔLRR.Sfu-Sty in vitro construct - deletion 0 Yes TlΔLRR.Sfu in vitro construct - deletion 0 Yes TlΔLRR.Sty in vitro construct - deletion 0 Yes TlΔLRR.T:Zzzz\FLAG in vitro construct - coding region fusion 0 Yes TlΔLRR.Xba.A200 in vitro construct 0 Yes TlΔLRR.Xba.A50 in vitro construct 0 Yes TlΔLRR.Xba in vitro construct - deletion 0 Yes TlΔLRR in vitro construct - deletion in vitro construct 0 Yes TlΔN2.Act5C in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes TlΔN3.Act5C in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes TlΔN4.Act5C.T:SV5\V5 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes TlΔN4.Act5C in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes TlΔN4.Scer\UAS.T:SV5\V5 in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes TlΔN5.Act5C in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes TlΔN6.Act5C in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes TlΔN6.C34.892A.Scer\UAS.T:SV5\V5 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlΔN6.C34A.Scer\UAS.T:SV5\V5 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlΔN6.C892A.Scer\UAS.T:SV5\V5 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes TlΔN6.Scer\UAS.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes TlΔN6.Scer\UAS.T:SV5\V5 in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes TlΔN6.SS.Scer\UAS.T:SV5\V5 in vitro construct - deletion 0 Yes TlΔN6.SSC892A.Scer\UAS.T:SV5\V5 in vitro construct - deletion in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes
Aneuploid Aberrations
Disrupted in	Df(3R)ME61 (Cook and Carpenter, 2002, Cook, 2000.10.9) Df(3R)ro80b (Cook and Carpenter, 2002, Rayburn et al., 2003) Df(3R)ro-XB3 (Rose et al., 1997, Anderson et al., 1985, Lieberfarb et al., 1996, Wu and Anderson, 1998) Df(3R)Tl-D (Anderson et al., 1985) Df(3R)Tl-I (Cook, 2000.10.9, Anderson et al., 1985, Cook and Carpenter, 2002) Df(3R)Tl-K (Anderson et al., 1985) Df(3R)Tl-P (Cook and Carpenter, 2002, Cook, 2000.10.9, Anderson et al., 1985) Df(3R)Tl-Q (Anderson et al., 1985) Df(3R)Tl-X (Rose et al., 1997, Cook and Carpenter, 2002, Rayburn et al., 2003, Lieberfarb et al., 1996, Anderson et al., 1985, Cook, 2000.10.9, Wu and Anderson, 1998) In(3LR)Tlrv18 (Anderson et al., 1985) In(3R)Tlrv5 (Anderson et al., 1985) In(3R)Tlrv6 (Anderson et al., 1985) In(3R)Tlrv12 (Anderson et al., 1985) In(3R)Tlrv17 (Anderson et al., 1985) In(3R)Tlrv19 (Anderson et al., 1985) T(2;3)Tlrv3 (Anderson et al., 1985) Tp(3;3)ME61 (Cook, 2000.10.9)
Not disrupted in	Df(3R)Espl3 (Cook and Carpenter, 2002)
Duplicated in	Ts(YLt;3Lt)A121+Ts(YSt;3Rt)B158 (Anderson et al., 1985)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data reporter construct P{1.2dR} No P{1.4RR(-)} No P{1.4RR} No P{1.05Rd} No P{1.7RS} No P{1.15Rd} No P{3X950Tl4} No P{4xCME-lacZ} No P{320Rd} No P{450dR} No P{600Rd} No P{620dR} No P{750dR} No P{770Rd} No P{900dR} No P{950Tl.mut1-4} No P{950Tl.mut1} No P{950Tl.mut3} No P{950Tl.mut4} No P{950Tl} No P{Tl/lacZ.6.5LR} No P{Tl/lacZ.6.5RL} No UAS construct P{GD305} NA P{KK103505} NA P{TRiP.GL00474} NA P{TRiP.JF01276} NA P{TRiP.JF01491} NA P{UAS-Tl.10b-3559} NA P{UAS-Tl.10b.FLAG} NA P{UAS-Tl.10b.Myc} NA P{UAS-Tl.10b.Y} NA P{UAS-Tl.10b} NA P{UAS-Tl.10b-Nae} NA P{UAS-Tl.10b-Nar} NA P{UAS-Tl.10b-Stu} NA P{UAS-Tl.10b-ΔPEST} NA P{UAS-Tl.C755Y.C781Y.C799Y} NA P{UAS-Tl.C755Y.C781Y} NA P{UAS-Tl.C781S} NA P{UAS-Tl.C781Y} NA P{UAS-Tl.FLAG} NA P{UAS-Tl.H} NA P{UAS-Tl.M} NA P{UAS-Tl.Myc} NA heat-shock construct P{Hsp83-Toll10b} NA characterization construct P{1.4-Toll} NA P{6.5-Toll} NA P{bcd-Toll10b} NA P{Mhc'-Tl} NA
Insertions
	Type of insertions Name Expression data insertion of mobile activating element P{EP}TlEP1051 NA miscellaneous insertions Mi{MIC}TlMI00181 NA Mi{MIC}TlMI01254 NA P{}Tlrv22 NA P{GT1}BG02320BG02320 NA P{SUPor-P}TlKG03609 NA P{UASp-YFP.Rab10.T23N}25a NA PBac{RB}Tle03133 NA insertion of enhancer trap binary system P{GawB}TlNP0756 No insertion of enhancer trap P{FZ}TlF336 No P{lwB}TlAK80 No P{PZ}TlrK344 No
Gene Ontology: Function, Process & Cellular Component ( 34 unique terms )
Terms Based on Experimental Evidence ( 17 terms )
Molecular Function
CV term Evidence References
protein binding inferred from physical interaction with spz (Gangloff et al., 2008) inferred from physical interaction with wek (Chen et al., 2006) TIR domain binding inferred from physical interaction with Myd88 (Tauszig-Delamasure et al., 2002)
Biological Process
CV term Evidence References
antifungal humoral response inferred from mutant phenotype (Lemaitre et al., 1996) antimicrobial humoral response inferred from genetic interaction with imd (Lemaitre et al., 1996) cytokinesis inferred from mutant phenotype (Kiger et al., 2003) dorsal/ventral axis specification inferred from mutant phenotype (Chen et al., 2006) heart development inferred from mutant phenotype (Wang et al., 2005) hemocyte proliferation inferred from mutant phenotype (Huang et al., 2005) innate immune response inferred from direct assay (Chamilos et al., 2008) positive regulation of antibacterial peptide biosynthetic process inferred from mutant phenotype (Lemaitre et al., 1997) positive regulation of antifungal peptide biosynthetic process inferred from mutant phenotype (Manfruelli et al., 1999, Lemaitre et al., 1997) regulation of hemocyte differentiation inferred from mutant phenotype (Zettervall et al., 2004) synapse assembly inferred from mutant phenotype (Suzuki et al., 2000) synaptic target inhibition inferred from mutant phenotype (Inaki et al., 2010)
Cellular Component
CV term Evidence References
ESCRT-0 complex inferred from direct assay (Huang et al., 2010) plasma membrane inferred from direct assay (Huang et al., 2010) colocalizes_with plasma membrane inferred from direct assay (Mavrakis et al., 2009) protein complex inferred from physical interaction with spz (Gangloff et al., 2008)
Terms Based on Predictions or Assertions ( 20 terms )
Molecular Function
CV term Evidence References
cytokine binding traceable author statement (Underhill, 2003) transmembrane signaling receptor activity inferred from electronic annotation with InterPro:IPR004075 (FlyBase Curators et al., 2004-) non-traceable author statement (van Eeden and St. Johnston, 1999, Lemaitre, 2000.12.20)
Biological Process
CV term Evidence References
defense response non-traceable author statement (Lemaitre, 2000.12.20) traceable author statement (Meister et al., 2000) defense response to fungus traceable author statement (Underhill, 2003, Janssens and Beyaert, 2003, Brennan and Anderson, 2004, Leclerc and Reichhart, 2004) defense response to Gram-positive bacterium traceable author statement (Underhill, 2003, Janssens and Beyaert, 2003, Leclerc and Reichhart, 2004) dorsal/ventral axis specification non-traceable author statement (Lall and Patel, 2001) traceable author statement (Meister et al., 2000, Beutler and Poltorak, 2001) embryonic pattern specification traceable author statement (Kimbrell and Beutler, 2001) hemocyte proliferation traceable author statement (Evans et al., 2003) hemopoiesis traceable author statement (Meister et al., 2000) immune response traceable author statement (Martin-Blanco, 2000, Schneider, 2000, Beutler and Poltorak, 2001) innate immune response non-traceable author statement (Kwiatkowski, 2005) traceable author statement (Underhill, 2003, Janssens and Beyaert, 2003) regulation of melanization defense response traceable author statement (Kurz and Ewbank, 2003) response to bacterium non-traceable author statement (Silverman, 2003) response to fungus traceable author statement (Medzhitov and Janeway, 2000, Kimbrell and Beutler, 2001) signal transduction inferred from electronic annotation with InterPro:IPR000157, InterPro:IPR004075 (FlyBase Curators et al., 2004-) Toll signaling pathway non-traceable author statement (van Eeden and St. Johnston, 1999, Lemaitre, 2000.12.20) zygotic specification of dorsal/ventral axis non-traceable author statement (Leclerc and Reichhart, 2004)
Cellular Component
CV term Evidence References
integral to membrane non-traceable author statement (Lemaitre, 2000.12.20) integral to plasma membrane non-traceable author statement (Swiss-Prot Project Members, 1988.11.1) intrinsic to membrane inferred from electronic annotation with InterPro:IPR004075 (FlyBase Curators et al., 2004-)
Sequence Ontology: Class of Gene
	gene   nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
Protein-protein
Interacting group Assay References
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions Tl allele Gene References EP1051EP1051 pnr (Pena-Rangel et al., 2002) Tl3 Myd88 (Charatsi et al., 2003) Tl4 cact (Belvin et al., 1995) Tl8 Stat92E (Remillieux-Leschelle et al., 2002) TlMhc.PR Fas3 (Rose and Chiba, 1999) Tlr1 ea (Chasan et al., 1992) Tlr3 mxc (Remillieux-Leschelle et al., 2002) Rel (De Gregorio et al., 2002) Tlr5 ea (Chasan et al., 1992) Tlrv1 mxc (Remillieux-Leschelle et al., 2002) Rel (De Gregorio et al., 2002) Tlrv13 ea (Chasan et al., 1992) Tlrv19 ea (Chasan et al., 1992) unspecified Myd88 (Kambris et al., 2003) Npc1b (Bhattacharya and Steward, 2003) wek (Chen et al., 2006)
External Data
Linkouts	DroID - A comprehensive database of gene and protein interactions. • FBgn0262473
Orthologs
Genome-wide drosophilid orthologs	Dana\GF16456 Dere\GG11504 Dgri\GH14238 Dmoj\GI22147 Dpse\Tl Dsec\GM10345 Dsim\Tl Dvir\Tl Dwil\GK13544 Dyak\Tl
Curated drosophilid orthologs	Dafs\Tl (Bartolome, 2005) Dmir\Tl (Bartolome, 2005) Dpse\Tl   Drep\Tl (Ranz et al., 2001, Ranz et al., 2001) Dsim\Tl   Dvir\Tl   Dyak\Tl
Linkouts
Stocks & Reagents
Stocks Listed in FlyBase ( 22 )
Bloomington	30898 w*; P{UASp-Tl.Venus}4 30899 w*; P{UASp-Tl.Venus}1 30901 w*; P{UASp-Tl.PA-GFP}10 30900 w*; P{UASp-Tl.PA-GFP}3/CyO 3238 Tlr3 ca1/TM3, Sb1 31044 y1 v1; P{TRiP.JF01491}attP2 31477 y1 v1; P{TRiP.JF01276}attP2 30652 y1 w*; Mi{MIC}TlMI00181 35628 y1 sc* v1; P{TRiP.GL00474}attP2
Harvard	- PBac{RB}Tle03133
Kyoto	106934 Tlr3 ca1/TM3, Sb1
VDRC	v100078 P{KK103505}VIE-260B
	More stocks available...
Genomic Clones ( 1 )
	BACR24O14 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 98 )
	Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see GBrowse for alignment of the cDNAs and ESTs to the gene model.
cDNA Clones, Fully Sequenced
BDGP DGC clones	GH03720 RE13664 RE46574
Other clones	RT10576 RT11167
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	LD02201 LD02343 LD07108 LD20812 LD38708 RE04720 RE08531 RE13560 RE24617 RE27166 RE38783 RE45888 RE46595 RE71437 RE73449
Other clones	14240 16452 16547 19945 21027 28969 42419 51494 54246 75216 81106 81254 97486 105856 106977 108833 119600 162724 163424 180092 193921 214289.3 214319.20 214405.14 272923 CK01.60D.D3 CK01.65A.E2 CK01.73B.C12 CK01.91C.F5 CK01.93B.E8 CK01.109A.D7 CK01.111C.E6 Dmel_dig_012311_0613_3504 EK048911 EK053049 EK057833 EK071623 EK072852 EK093801 EK099216 EK116450 EK127925 EK136936 EK157356 EK158937 EK162438 EK166822 EK207258 EK214252 EK226506 EK230905 EK232912 EK235627 EK239536 EK244155 EK266949 EK267841 EK286901 EK297225 EK299744 EP04314 RP004027858 RP004069958 RP004081328 RP004157096 RP004203409 RP004220420 RP004239315 RP004248299 RP004248642 RP004250062 RP004256268 RP004393644 RP004422338 RP004441385 RP004536604 RP004537579 RP004537729
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0262473 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 43222
Antibody Information
	polyclonal (Gay and Keith, 1992)
Other Information
Discoverer
	Wieschaus and Nusslein-Volhard.
Etymology
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of	Source for merge of: Tl EP1051 (Cao et al., 2008)
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. (Armstrong et al., 1998)
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. (LeMosy, 2006) 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. (Kiger et al., 2003) 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) 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. (Weber et al., 2003) Tl regulates Drs expression through Dif. (Rutschmann et al., 2000) nec negatively regulates the Tl signalling pathway. (Levashina et al., 1999) Tl does not function as a pattern recognition receptor in the Drosophila host defence. (Levashina et al., 1999) Tl acts in a cell autonomous manner in the fat body. (Manfruelli et al., 1999) A linear activation cascade spz-Tl-cact-dl/Dif leads to the induction of the Drs gene in larval fat body cells. (Manfruelli et al., 1999) 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. (Halfon and Keshishian, 1998) Tl mRNA is translationally activated by regulated cytoplasmic polyadenylation. (Schisa and Strickland, 1998) Targeting of either tub or pll product to the plasma membrane by myristylation is sufficient to activate the signal transduction pathway that leads to translocation of the dl product. Activated Tl induces a localized recruitment of tub and pll proteins to the plasma membrane. (Towb et al., 1998) Tl pathway is required for the nuclear import of dl in the immune response, but not required for the nuclear import of Dif. Cytoplasmic retention of both dl and Dif depends on cact protein. The two signalling pathways that target cact for degradation must discriminate between cact-dl and cact-Dif complexes. (Wu and Anderson, 1998) 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. (Huang et al., 1997) 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. (Rose et al., 1997) 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. (Lemaitre et al., 1996) Tl and pll can functionally interact to enhance dl activity synergistically. (Norris and Manley, 1996) 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. (Belvin et al., 1995) 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. (Halfon et al., 1995) 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) dl is not involved in the formation of melanotic tumours of Tl mutations. (Lemaitre et al., 1995) tub is capable of acting as both a chaperon or escort for dl as it moves to the nucleus and then as a transcriptional coactivator. The intracytoplasmic domain of Tl is sufficient for activating the signalling pathway that leads to dl-tub nuclear translocation in Schneider cells. (Norris and Manley, 1995) Studies on a truncated Tl receptor indicate that the Tl receptor extracellular domain regulates the intrinsic signaling activity of its cytoplasmic domain. (Winans and Hashimoto, 1995) dl is an embryonic phosphoprotein and its phosphorylation state is regulated by an intracellular signaling pathway initiated by the transmembrane receptor Tl. Using a combined genetic and biochemical approach it is demonstrated that activation of Tl stimulates an increase in the extent of dl phosphorylation. (Gillespie and Wasserman, 1994) Dorsal-ventral patterning is regulated by a signalling pathway that includes Tl and transcription factors, dl, that interact with related enhancers, rho. The κ enhancer from mouse is capable of generating lateral stripes of Ecol\lacZ gene expression in transgenic embryos in a pattern similar to that directed by rho enhancer. Results suggest that enhancers can couple conserved signalling pathways to divergent gene functions, dorso-ventral patterning and mammalian haematopoiesis. (Gonzalez-Crespo and Levine, 1994) 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. (Morisato and Anderson, 1994) Comparisons of early development to that in other insects have revealed conservation of some aspects of development, as well as differences that may explain variations in early patterning events. (Patel, 1994) The Tl signalling pathway generates a dl nuclear gradient which initiates the differentiation of the mesoderm, neuroectoderm and dorsal ectoderm by activating and repressing gene expression in the early embryo. A second signalling pathway controlled by the tor receptor kinase also modulates dl activity. (Rusch and Levine, 1994) 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. (Wharton et al., 1994) Increased cytoplasmic calcium concentration and the expression of constitutively active Tl receptors can induce the relocalisation of dl in culture cells. Activation of endogenous Pka-C1, expression of wild type Tl receptors or treatment of cells with activators of Pkc53E and radical oxygen intermediates have only a marginal effects on the cellular distribution of dl protein. (Kubota et al., 1993) 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. (Roth, 1993) 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. (Ruohola-Baker et al., 1993) 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). (Wharton and Crews, 1993) 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. (Chasan et al., 1992) Toll enhances transport of dl protein into nucleus in cotransfected Schneider cells, perhaps via activated protein kinase A that phosphorylates dl gene product. (Norris and Manley, 1992) 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. (Stein and Nusslein-Volhard, 1992) The properties of a peptide corresponding to residues 166-188 of the Tl protein have been studied in vitro. (Gay et al., 1991) The cytoplasmic domain of the Tl protein is related to that of the human interleukin-1 receptor. (Gay and Keith, 1991) The Tl protein is a glycoprotein which is tightly associated with embryonic membranes. (Hashimoto et al., 1991) Recessive dorsalizing mutants of the dorsal group gene Tl have significantly reduced axial ratios in pupae. (Letsou et al., 1991) Mutations in maternal dorsal class gene Tl do not interact with RpII140wimp. (Parkhurst and Ish-Horowicz, 1991) Local activation of Tl by a Tl ligand initiates the formation of the dl nuclear concentration gradient, thereby determining the dorsoventral pattern. (Stein et al., 1991) 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. (Yasuda et al., 1991) sim gene product is required for the normal expression of Tl. (Nambu et al., 1990) 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. (Rushlow and Levine, 1990) Genetic and molecular analysis demonstrates that Tl is expressed and is functional zygotically as well as maternally. (Gerttula et al., 1988) Epistatic relationships exist between dorsalizing maternal effect mutations and "dppHin" alleles. (Irish and Gelbart, 1987) The expression of genes controlling neurogenesis is dependent on the previous activity of the genes controlling the development of the embryonic dorsal-ventral pattern. 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. (Campos-Ortega, 1983) 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).
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_079794 NM_170287 RefSeq (Proteins) • NP_524518 NP_733166 DDBJ / EMBL / GenBank • AAA28941 AAF56624 AAM51943 AAN14086 AAQ64932 AAQ64933 AAQ64934 AAQ64935 AAQ64936 AAQ64937 AAQ64938 AI063746 AQ025161 AQ026361 AW940139 AY121616 AY349649 AY349650 AY349651 AY349652 AY349653 AY349654 AY349655 BI362102 CZ480900 M19969 Entrez Gene - A searchable database of RefSeq genes. • 43222 UniProtKB/Swiss-Prot • P08953
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Toll/interleukin-1 receptor homology (TIR) domain (IPR000157) Leucine-rich repeat-containing N-terminal (IPR000372) Cysteine-rich flanking region, C-terminal (IPR000483) Leucine-rich repeat (IPR001611) Leucine-rich repeat, typical subtype (IPR003591) Interleukin-1 receptor, type I/Toll precursor (IPR004075)
Linkouts
	DroID - A comprehensive database of gene and protein interactions. • FBgn0262473 DRSC - Results from RNAi screens. • FBgn0262473 FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0262473 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 43222 modMine - Data generated by the modENCODE project. • FBgn0262473
Synonyms & Secondary IDs ( 23 )
Reported As
Symbol Synonym	CG5490 (Asha et al., 2003, De Gregorio et al., 2002, De Gregorio et al., 2001, Pal et al., 2007, Keleman et al., 2009.8.5, Hernandez-Romano et al., 2008, Wertheim et al., 2005, Ni et al., 2008, Ni et al., 2011.7.19) CT17414 (Hynes and Zhao, 2000) dToll (da Silva Correia et al., 2001, Aderem and Ulevitch, 2000) dToll1 (Singh et al., 2003) EP1051 (Cao et al., 2008, ) EP(3)1051 (Pena-Rangel et al., 2002) Fs(1)Tl (Campos-Ortega, 1983) Fs(3)Tl   mat(3)9   mel(3)9   mel(3)10   T1 (Asha et al., 2003, Bartolome and Charlesworth, 2006) tl (Morrison et al., 2000) Tl (Castillejo-Lopez and Haecker, 2005, Shen and Tanda, 2007, Araujo and Bier, 2000, Carneiro et al., 2006, LeMosy, 2006, Kambris et al., 2006, Senger et al., 2006, Kim et al., 2006, Mizutani et al., 2005, Brun et al., 2006, Mizutani et al., 2006, Zhang et al., 2007, Taylor and Kimbrell, 2007, Cowden and Levine, 2003, Wang et al., 2005, Xing et al., 2007, Zeitouni et al., 2007, Minidorff et al., 2007, Markstein et al., 2008, Tan et al., 2008, Gilchrist et al., 2008, Wu et al., 2007, Jin et al., 2009, Buchon et al., 2009, Witzberger et al., 2008, Lamaris et al., 2009, Leulier et al., 2003, Chamilos et al., 2008, Chamilos et al., 2009, Thoetkiattikul et al., 2005, Zeitlinger et al., 2007, Buchon et al., 2009, Zhu et al., 2008, Ahmad et al., 2009, Obbard et al., 2009, Tauszig-Delamasure et al., 2002, Keranen et al., 2006, Morozova et al., 2007, Wertheim et al., 2005, Goto et al., 2010, Vonkavaara et al., 2008, Inaki et al., 2010, Valanne et al., 2010, Chen et al., 2010, Kong et al., 2010, Wang et al., 2011, Stein et al., 2010, Kim et al., 2010) TL (Luo et al., 2001) Toll (Diangelo et al., 2009, Quintin et al., 2007, Sandmann et al., 2007, Goto et al., 2003, Kuranaga and Miura, 2007, Vogler et al., 2008, Reichhart, 2008, Weber et al., 2007, Gangloff et al., 2008, Staudt et al., 2005, Hendrix et al., 2008, Korolchuk et al., 2007, Liberman et al., 2009, Shia et al., 2009, Wagner et al., 2008, Blanco and Gehring, 2008) toll (Janssens and Beyaert, 2002, Vogel, 1998) Toll-1 (Mushegian and Medzhitov, 2001, Medzhitov, 2001) toll-1 (Janssens and Beyaert, 2002) Toll1 (Shia et al., 2009)
Name Synonym	EP1051   Protein toll precursor (Hernandez-Romano et al., 2008) Toll (Chen et al., 2010, Clark et al., 2011, Galac and Lazzaro, 2011, Cao et al., 2008, Cui et al., 2008, Furlong et al., 2001, Meng et al., 1999, Braun et al., 1998, Uttenweiler-Joseph et al., 1998, Govind et al., 1998, Ntwasa et al., 1997, Ferrandon et al., 1997, Schisa and Strickland, 1996, Rosetto et al., 1995, Wharton and Crews, 1993, Leptin et al., 1992, Schupbach and Wieschaus, 1986, Kambris et al., 2006, Mace et al., 2005, Pal et al., 2007, Zhou et al., 2005, Pal and Wu, 2005, Minakhina and Steward, 2006, Kuttenkeuler and Boutros, 2007, Tao et al., 2007, Pham et al., 2007, Araujo et al., 2006, Scherfer et al., 2006, Araujo and Bier, 2000, Akira et al., 2006, Chen et al., 2006, Chen, 2006, Biemar et al., 2006, LeMosy, 2006, Sambandan et al., 2006, Mulinari et al., 2006, Fenckova and Dolezal, 2007, Scherfer et al., 2007, Zeitlinger et al., 2007, Zaffran et al., 2006, Ulvila et al., 2006, Jang et al., 2006, Brun et al., 2006, Kleino et al., 2008, Tao et al., 2007, Waterhouse et al., 2007, Rutschmann et al., 2000, Guan et al., 2006, Park et al., 2003, Kuranaga and Miura, 2007, Araujo et al., 2008, DiAngelo et al., 2008, Tanda et al., 2008, Taylor and Kimbrell, 2007, Christophides et al., 2002, Davidson and Erwin, 2006, Goto et al., 2008, Cowden and Levine, 2003, Ritzenthaler and Chiba, 2003, Stein et al., 2008, Staudt et al., 2005, Wang et al., 2005, Tanji et al., 2007, Xing et al., 2007, Tao et al., 2007, Zeitouni et al., 2007, Busse et al., 2007, Qi et al., 2008, Inaki et al., 2007, Tan et al., 2008, Wu and Sato, 2008, Zhu et al., 2008, Liu et al., 2009, Mavrakis et al., 2009, Wu et al., 2007, Jin et al., 2009, Buchon et al., 2009, Witzberger et al., 2008, Ooi et al., 2002, Leulier et al., 2003, Roxstrom-Lindquist et al., 2004, Gesellchen et al., 2005, Stramer et al., 2008, Luo et al., 2001, Chamilos et al., 2009, Ahmad et al., 2009, Cronin et al., 2009, Zeitlinger et al., 2007, Shi et al., 2006, Zeitlinger et al., 2007, Buchon et al., 2009, Tauszig-Delamasure et al., 2002, Keranen et al., 2006, Morozova et al., 2007, Zimmermann et al., 2006, Coll et al., 2010, Huang et al., 2010, Vonkavaara et al., 2008, Tanji et al., 2010, Inaki et al., 2010, Junell et al., 2010, Arnot et al., 2010, Valanne et al., 2010, Yagi and Ip, 2005, Zsindely et al., 2009, Wang et al., 2011, Stein et al., 2010, Fulkerson and Estes, 2011, Gordon et al., 2008, Vazquez-Pianzola et al., 2011, Kim et al., 2010) toll (Muller et al., 2005, Boettiger and Levine, 2009) Toll-1 (Waterhouse et al., 2007, Ooi et al., 2002, Narbonne-Reveau et al., 2011) toll receptor (Haesemeyer M et al., 2007) Toll receptor (Tipping and Veraksa, 2008)
Secondary FlyBase IDs
	FBgn0003717 FBgn0062707
References ( 666 )
Generate a list of	All references relating to this gene ( 666 )
List References by type	Research paper  ( 349 ) Supplementary material  ( 11 ) Review  ( 166 ) Personal communication to FlyBase  ( 8 ) Abstract  ( 110 ) FlyBase analysis  ( 3 ) Stock list  ( 1 ) Letter  ( 3 ) Protein sequence record  ( 1 ) Book  ( 1 ) Computer file  ( 4 ) Conference report  ( 2 ) Automatic genome annotation  ( 1 ) Book review  ( 2 ) Interview  ( 1 ) Poem  ( 1 ) Curated genome annotation  ( 1 ) Biography  ( 1 )
Recent research papers ( 20 )
	Clark et al., 2011, Curr. Biol. 21(19): 1672--1677 Multiple TGF-β Superfamily Signals Modulate the Adult Drosophila Immune Response. [FBrf0216429] Fulkerson and Estes, 2011, J. Exp. Zool. B Mol. Dev. Evol. 316B(1): 61--75 Common motifs shared by conserved enhancers of Drosophila midline glial genes. [FBrf0212527] Galac and Lazzaro, 2011, Microbes Infect. 13(7): 673--683 Comparative pathology of bacteria in the genus Providencia to a natural host, Drosophila melanogaster. [FBrf0214338] Herren and Lemaitre, 2011, Cell. Microbiol. 13(9): 1385--1396 Spiroplasma and host immunity: activation of humoral immune responses increases endosymbiont load and susceptibility to certain Gram-negative bacterial pathogens in Drosophila melanogaster. [FBrf0214693] Morozova et al., 2011, Genetics 187(4): 1193--1205 Transcriptional networks for alcohol sensitivity in Drosophila melanogaster. [FBrf0214377] Narbonne-Reveau et al., 2011, PLoS ONE 6(2): e17470 Lack of an antibacterial response defect in Drosophila toll-9 mutant. [FBrf0213211] Vazquez-Pianzola et al., 2011, Dev. Biol. 357(2): 404--418 Pabp binds to the osk 3'UTR and specifically contributes to osk mRNA stability and oocyte accumulation. [FBrf0214798] Wang et al., 2011, Infect. Immun. 79(2): 606--616 Host and Pathogen Glycosaminoglycan-Binding Proteins Modulate Antimicrobial Peptide Responses in Drosophila melanogaster. [FBrf0212806] Arnot et al., 2010, J. Biol. Chem. 285(25): 19502--19509 Molecular mechanism that induces activation of spatzle, the ligand for the Drosophila toll receptor. [FBrf0211038] Chen et al., 2010, Proc. Natl. Acad. Sci. U.S.A. 107(48): 20774--20779 Participation of the p38 pathway in Drosophila host defense against pathogenic bacteria and fungi. [FBrf0212390] Coll et al., 2010, Genes Dev. 24(2): 129--134 A novel, noncanonical mechanism of cytoplasmic polyadenylation operates in Drosophila embryogenesis. [FBrf0209762] Goto et al., 2010, J. Biol. Chem. 285(21): 15731--15738 Cooperative regulation of the induction of the novel antibacterial Listericin by peptidoglycan recognition protein LE and the JAK-STAT pathway. [FBrf0210770] Huang et al., 2010, Proc. Natl. Acad. Sci. U.S.A. 107(18): 8322--8327 Endocytic pathway is required for Drosophila Toll innate immune signaling. [FBrf0210737] Inaki et al., 2010, Development 137(13): 2139--2146 Drosophila Tey represses transcription of the repulsive cue Toll and generates neuromuscular target specificity. [FBrf0210991] Junell et al., 2010, Mol. Cell. Biol. 30(14): 3672--3684 The POU transcription factor Drifter/Ventral veinless regulates expression of Drosophila immune defense genes. [FBrf0211117] Kim et al., 2010, Exp. Gerontol. 45(12): 984--987 Enhancement of stress resistances and downregulation of Imd pathway by lower developmental temperature in Drosophila melanogaster. [FBrf0212401] Kong et al., 2010, Alcoholism, Clin. Exp. Res. 34(2): 302--316 Ethanol-regulated genes that contribute to ethanol sensitivity and rapid tolerance in Drosophila. [FBrf0209720] Stein et al., 2010, Curr. Biol. 20(21): 1953--1958 Localization and Activation of the Drosophila Protease Easter Require the ER-Resident Saposin-like Protein Seele. [FBrf0212238] Tanji et al., 2010, Proc. Natl. Acad. Sci. U.S.A. 107(33): 14715--14720 Heterodimers of NF-{kappa}B transcription factors DIF and Relish regulate antimicrobial peptide genes in Drosophila. [FBrf0211616] Valanne et al., 2010, J. Immunol. 184(11): 6188--6198 Genome-wide RNA interference in Drosophila cells identifies G protein-coupled receptor kinase 2 as a conserved regulator of NF-kappaB signaling. [FBrf0210846] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...