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Gene Dmel\tll

General Information
Symbol	Dmel\tll	Species	D. melanogaster
Name	tailless	Annotation symbol	CG1378
Feature type	protein_coding_gene	FlyBase ID	FBgn0003720
Created / Updated	2005-01-10/2005-01-10
Genomic Location
Chromosome (arm)	3R	Recombination map	3-102
Cytogenetic map	100A6-100A6	Sequence location	3R:26,678,037..26,680,095 [+]
Map ( 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 tailless is referred to in FlyBase by the symbol tll (CG1378, FBgn0003720). It has the cytological map location 100A6. Its sequence location is 3R:26678037..26680095. Its molecular function is described as: ligand-dependent nuclear receptor activity; transcription factor activity; DNA binding; steroid hormone receptor activity; zinc ion binding; sequence-specific DNA binding. It is involved in the biological processes described with 12 unique terms, many of which group under: anatomical structure development; cell fate commitment; anterior/posterior axis specification; transcription, DNA-dependent; sensory organ development; cell differentiation; axis specification; organ development; RNA metabolic process; transmembrane receptor protein tyrosine kinase signaling pathway; regulation of metabolic process. 27 alleles are reported. The phenotypes of these alleles are annotated with 26 unique terms, many of which group under: organ system; anatomical structure; nervous system; larval head segment; embryonic nervous system; primordium; abdominal segment 8; embryonic abdomen; abdominal segment 7; endocrine system; ectoderm derivative; spiracle; cephalopharyngeal skeleton; peripheral nervous system; late extended germ band embryo. It has one annotated transcript and one annotated polypeptide.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	tll: tailless (J. A. Lengyel) Recessive; zygotic lethal with pattern deletions in anterior and posterior of embryo (but body of normal length due to increase in length of non-deleted pattern elements). Anteriorly, dorsal portion of cephalopharyngeal skeleton is defective (dorsal arms shortened, dorsal bridge unfused), dorsal pouch shortened and scleritized, much of brain missing. Posteriorly, eighth abdominal segment and telson, Malpighian tubules, hindgut and much of posterior midgut missing.
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 100A6-100A6   Limits computationally determined from genome sequence between P{lacW}l(3)s2500s2500 and P{PZ}Aph-407028&P{PZ}dcorK215
Experimentally Determined Cytological Location
Cytogenetic map Notes References 100A6-100B1 (determined by in situ hybridisation)   (Ranz et al., 1997) 100B1-100B2 (determined by in situ hybridisation)   (Pignoni et al., 1990)
Experimentally Determined Recombination Data
Location	3-102 (Tearle and Nusslein-Volhard, 1987)
Left of (cM)
Right of (cM)
Notes
Molecular Map Data
Gene Order (in direction of increasing cytology) References In direction of increasing cytology: tll? Ptx1+ (Vorbruggen et al., 1997) Gene Order (overall orientation not stated) References
Gene Model & Products
Please see the GBrowse view of Dmel\tll for information on other features
Comments on Gene Model
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) tll-RA FBtr0085709  NM_079857   1938   452
Additional Transcript Data & Comments
Reported size (kB)	2.0 (northern blot) (Pignoni et al., 1990)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kD) Length (aa) Theoretical pI RefSeq ID GenBank protein tll-PA   FBpp0085071   50.5   452   9.10     NP_524596   AAF57091
Additional Polypeptide Data & Comments
Reported size (kD)	452 (aa); 50.5 (kD) (Pignoni et al., 1990)
Comments
External Data
Linkouts	PANTHER - Protein classification by function, families, and pathways • FBgn0003720
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Zinc finger, NHR/GATA-type (IPR013088) Vitamin D receptor (IPR000324) Nuclear hormone receptor, ligand-binding, core (IPR000536) Zinc finger, nuclear hormone receptor-type (IPR001628) Steroid hormone receptor (IPR001723) Nuclear hormone receptor, ligand-binding (IPR008946) TRANSFAC - Eukaryotic transcription factors, their genomic binding sites, and DNA-binding profiles • T00789
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AE003775 AAF57091   AF019362 AAB71371   BT022195   M34639 AAA28936
UniProtKB/Swiss-Prot	P18102
UniProtKB/TrEMBL
Maps to
Does NOT map to
Identified with	IP01133 (Hoskins et al., 2005) D1195 (BDGP Project Members, 1994-1999)
Mapped Features & Mutations
Please see GBrowse or insertion reports for information on insertions of transgenic constructs and features not listed here Type Symbol & Location Additional Notes References aberration junction In(3R)tll.bk1 3R:26,677,122..26,678,031 evidence=experimental linked_to=SstI-PstI_rfrag (Pignoni et al., 1990) point mutation tll[1] 3R:26,678,490..26,678,490 evidence=experimental na_change=G26678490A pr_change=C34Y|tll-PA reported_na_change=G454A reported_pr_change=C34Y (Diaz et al., 1996) point mutation tll[le3] 3R:26,678,516..26,678,516 evidence=experimental na_change=G26678516A pr_change=G43S|tll-PA reported_na_change=G480A reported_pr_change=G43S (Diaz et al., 1996) point mutation tll[l29] 3R:26,678,597..26,678,597 evidence=experimental na_change=T26678597C pr_change=C70R|tll-PA reported_na_change=T561C reported_pr_change=C70R (Diaz et al., 1996) point mutation tll[l49] 3R:26,678,623..26,678,623 evidence=experimental na_change=T26678623A pr_change=C78@|tll-PA reported_na_change=T596A reported_pr_change=C78@ (Diaz et al., 1996) protein binding site tll-protein_bind-1 3R:26,676,912..26,676,926 bound_moiety=bcd-XP comment=High affinity bcd binding site evidence=experimental (Liaw and Lengyel, 1993, Bergman et al., 2004) protein binding site tll-protein_bind-10 3R:26,677,882..26,677,890 bound_moiety=ttk-XP evidence=experimental (Chen et al., 2002, Bergman et al., 2004) protein binding site tll-protein_bind-11 3R:26,677,844..26,677,856 bound_moiety=Trl-XP evidence=experimental (Liaw et al., 1995, Bergman et al., 2004) protein binding site tll-protein_bind-12 3R:26,677,189..26,677,208 bound_moiety=bcd-XP evidence=experimental (Liaw and Lengyel, 1993, Bergman et al., 2004) protein binding site tll-protein_bind-13 3R:26,677,844..26,677,873 evidence=experimental (Liaw et al., 1993, Bergman et al., 2004) protein binding site tll-protein_bind-14 3R:26,677,755..26,677,774 evidence=experimental (Bergman et al., 2004, Liaw et al., 1993) protein binding site tll-protein_bind-15 3R:26,676,939..26,676,953 bound_moiety=bcd-XP evidence=experimental (Bergman et al., 2004, Liaw and Lengyel, 1993) protein binding site tll-protein_bind-16 3R:26,677,900..26,677,936 evidence=experimental (Liaw et al., 1993, Bergman et al., 2004) protein binding site tll-protein_bind-2 3R:26,677,067..26,677,081 bound_moiety=bcd-XP evidence=experimental (Liaw and Lengyel, 1993, Bergman et al., 2004) protein binding site tll-protein_bind-3 3R:26,676,864..26,676,878 bound_moiety=bcd-XP evidence=experimental (Bergman et al., 2004, Liaw and Lengyel, 1993) protein binding site tll-protein_bind-4 3R:26,677,842..26,677,864 bound_moiety=grh-XP evidence=experimental (Liaw et al., 1995, Bergman et al., 2004) protein binding site tll-protein_bind-5 3R:26,676,842..26,676,856 bound_moiety=bcd-XP evidence=experimental (Liaw and Lengyel, 1993, Bergman et al., 2004) protein binding site tll-protein_bind-6 3R:26,677,806..26,677,837 evidence=experimental (Bergman et al., 2004, Liaw et al., 1993) protein binding site tll-protein_bind-7 3R:26,676,964..26,676,978 bound_moiety=bcd-XP evidence=experimental (Bergman et al., 2004, Liaw and Lengyel, 1993) protein binding site tll-protein_bind-8 3R:26,677,706..26,677,739 evidence=experimental (Liaw et al., 1993, Bergman et al., 2004) protein binding site tll-protein_bind-9 3R:26,677,097..26,677,125 bound_moiety=bcd-XP evidence=experimental (Bergman et al., 2004, Liaw and Lengyel, 1993) rescue fragment tll[+t10.2] 3R:26,671,815..26,682,115 comment=Approximate position of rescue fragment which includes about 6.2kb upstream and 2kb downstream of the transcription unit. evidence=experimental (Pignoni et al., 1990)
External Data
Linkouts	DEDB - Drosophila exon database: splicing graphs • 11408
Crossreferences
Expression Data
FlyBase-Curated Data
Transcript and Protein data	Please see the FlyBase Gene Expression Report for details of gene expression from the literature.
Summary of Transcript Expression
Stage Tissue/Position Reference  embryonic stage | stage 7,8  Pc3 neuroblasts (Rudolph et al., 1997)  embryonic stage  procephalic neuroblasts (Rudolph et al., 1997)  embryonic stage | syncytial blastoderm   (Pignoni et al., 1992)  embryonic stage | stage 12  optic lobe | primoridum (Rudolph et al., 1997)  embryonic stage | cellular blastoderm   (Pignoni et al., 1992)  embryonic stage | syncytial blastoderm   (Pignoni et al., 1990)  embryonic stage | stage 7,8  Pc2 neuroblasts (Rudolph et al., 1997)  embryonic stage | syncytial blastoderm   (Pignoni et al., 1992)  embryonic stage | stage 15  visual primordium (Li et al., 2002)  embryonic stage | stage 5   (Wu and Lengyel, 1998)  embryonic stage  embryonic protocerebral neuromere (Rudolph et al., 1997)  embryonic stage | stage 10,11  posterior protocerebral neuroblasts (Rudolph et al., 1997)  embryonic stage | stage 10  intercalary segment (Li et al., 2002)  embryonic stage | stage 5   (Wu and Lengyel, 1998)  embryonic stage | stage 5   (Wu and Lengyel, 1998)  embryonic stage | stage 15  posterior deutocerebral neuromere (Li et al., 2002)  embryonic stage | syncytial blastoderm   (Pignoni et al., 1990)  embryonic stage | cellular blastoderm   (Pignoni et al., 1992)  embryonic stage  larval brain (Pignoni et al., 1990)  embryonic stage | stage 9,10  Pa4 neuroblasts (Rudolph et al., 1997)  embryonic stage | stage 5   (Wu and Lengyel, 1998)  embryonic stage | cellular blastoderm   (Pignoni et al., 1990)  embryonic stage | stage 9,10  Pa3 neuroblasts (Rudolph et al., 1997)  embryonic stage | stage 15  protocerebrum (Li et al., 2002)  embryonic stage | cellular blastoderm   (Pignoni et al., 1990)  embryonic stage | early   (Perkins et al., 1992)  embryonic stage | stage 10  procephalic neurectoderm (Li et al., 2002)  embryonic stage | stage >=4  procephalic neurectoderm (Rudolph et al., 1997)  embryonic stage,larval stage  optic lobe (Rudolph et al., 1997)
Marker for
Subcellular Localization
CV Term
Summary of Polypeptide Expression
Stage Tissue/Position Reference  embryonic stage | syncytial blastoderm   (Pignoni et al., 1992)  embryonic stage | syncytial blastoderm   (Pignoni et al., 1992)  embryonic stage | cellular blastoderm   (Pignoni et al., 1992)  embryonic stage | cellular blastoderm   (Pignoni et al., 1992)
Marker for
Subcellular Localization
CV Term
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003720 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • FBgn0003720 GEO (NCBI) - Gene expression data: microarray and other high-throughput technologies • FBgn0003720
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal tllhs.PS tlll49 lethal | recessive tll10 tll11 tll1 tll3 tll4 tll5 tll6 tll7 tll8 tll9 lethal | embryonic stage | recessive tll1 tll2 tlll29 tlll49 lethal | embryonic stage | partially | recessive tllle3 lethal | larval stage tll2 Other Phenotypes Allele wild-type tll+t10.2 Phenotype manifest in Allele cephalopharyngeal skeleton tll1 filzkorper tll1 tlll29 larval anus & embryo tll1 embryonic/larval posterior spiracle & embryo tll1 larval hindgut & embryo tll1 tll2 tlll29 tlll49 tllle3 Malpighian tubule tll1 longitudinal visceral muscle primordium tll1 abdominal segment 8 tll1 tlla tllc tlle tlll49 tllunspecified embryonic abdominal segment 8 tll1 anal pad tll1 tlll29 tllle3 Malpighian tubule primordium tll1 abdominal segment 2 tll1 abdominal segment 3 tll1 abdominal segment 4 tll1 abdominal segment 5 tll1 abdominal segment 6 tll1 abdominal segment 7 tll1 telson tll1 tll2 tlla tllc tlle tllunspecified tracheal pit tll1 gonad tllL1 embryonic central nervous system tlla Bolwig's organ tllhs.PS tlll49 larval optic lobe tllhs.PS tlll49 embryonic dorsal epidermis tllhs.PS denticle belt & abdominal segment 8 tlll29 anterior protocerebral neuromere tlll49 posterior protocerebral neuromere tlll49 embryonic brain tlll49 embryonic corpus cardiacum tlll49 embryo | gap tllunspecified
Classical Alleles ( 21 )
For All Classical Alleles Show
Allele of tll Class Mutagen Stocks Known lesion tll10 loss of function ethyl methanesulfonate 0 -- tll11 loss of function ethyl methanesulfonate 0 -- tll1 loss of function, hypomorph ethyl methanesulfonate 6 Yes tll2 hypomorph spontaneous X ray 0 Yes tll3 loss of function ethyl methanesulfonate 0 -- tll4 loss of function ethyl methanesulfonate 0 -- tll5 loss of function ethyl methanesulfonate 0 -- tll6 loss of function ethyl methanesulfonate 0 -- tll7 loss of function ethyl methanesulfonate 0 -- tll8 loss of function ethyl methanesulfonate 0 -- tll9 loss of function ethyl methanesulfonate 0 -- tllG 0 -- tllL1 0 -- tlla hypomorph X ray 0 -- tllc X ray 0 -- tlle 0 -- tllg loss of function 0 -- tlll29 X ray 0 Yes tlll49 X ray 1 Yes tllle3 ethyl methanesulfonate 0 Yes tllunspecified ethyl methanesulfonate 0 --
Alleles Carried on Transgenic Constructs ( 6 )
For All Alleles Carried on Transgenic Constructs Show
Allele of tll Class Mutagen Stocks Known lesion tll+t10.2 in vitro construct | genomic fragment 0 Yes tllGD1548 in vitro construct | RNAi in vitro construct | regulatory fusion 1 Yes tllScer\UAS.cHa in vitro construct | regulatory fusion 0 Yes tllhs.PH in vitro construct | regulatory fusion 0 Yes tllhs.PS in vitro construct | regulatory fusion 0 Yes tllhs.T:Scer\GAL4.T:Ivir\HA1 in vitro construct | regulatory fusion in vitro construct | coding region fusion 0 Yes
Aneuploid Aberrations
Useful deficiency	Df(3R)tll-e
Useful duplication	Dp(3;3)516
Disrupted in	Df(3R)A113 (Zilian et al., 1999, Jurgens et al., 1984, Strecker et al., 1986, Justice et al., 1995) Df(3R)A177.X1 (Zilian et al., 1999, Justice et al., 1995) Df(3R)A177.X3 (Zilian et al., 1999) Df(3R)A177der20 (Zilian et al., 1999, Jursnich et al., 1990, Justice et al., 1995) Df(3R)A177der21 (Zilian et al., 1999, Justice et al., 1995) Df(3R)A177der25 (Zilian et al., 1999, Justice et al., 1995) Df(3R)A177der26 (Zilian et al., 1999, Justice et al., 1995) Df(3R)Exel6217 (Ryder, 2004.4.26) Df(3R)Exel8194 (Ryder, 2004.4.26) Df(3R)J55 (Jurgens et al., 1984, Strecker et al., 1988) Df(3R)PH3 (Zilian et al., 1999) Df(3R)R97 (Strecker et al., 1988) Df(3R)tll-e (Cook, 2006.1.31, Justice et al., 1995) Df(3R)tll-g (Cook, 2006.1.31, Harbecke and Lengyel, 1995, Klingler et al., 1996) Df(3R)tll-pgx (Margolis et al., 1995, Diaz et al., 1996) Dp(3;1)150P (Strecker et al., 1988, Pignoni et al., 1990) Dp(3;1)34 (Strecker et al., 1988) In(3R)C (Tearle and Nusslein-Volhard, 1987) In(3R)tll (Strecker et al., 1986, Diaz et al., 1996) Ts(YSt;3Lt)A113+Dp(3;1)150P (Jursnich et al., 1990) Ts(YSt;3Lt)G75 (Jurgens et al., 1984)
Not disrupted in	Df(3R)A177der22 (Zilian et al., 1999) Df(3R)P75A.A1 (Justice et al., 1995) Df(3R)PGX8 (Zilian et al., 1999) Df(3R)X3F (Cook, 2006.1.31) Dp(3;1)1A (Strecker et al., 1988) Dp(3;1)48 (Strecker et al., 1988) Dp(3;1)79 (Strecker et al., 1988) Dp(3;1)93 (Strecker et al., 1988)
Duplicated in	Ts(YLt;3Rt)A113 (Justice et al., 1995) Ts(YSt;3Rt)B226 (Jurgens and Weigel, 1988) Ts(YSt;3Rt)J116 (Jurgens and Weigel, 1988)
Not duplicated in	Dp(3;1)Dr-RXT11 (Jurgens and Weigel, 1988)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data reporter construct P{G11} No P{G13} No P{G14} No P{G15} No P{G16} No P{G17} No P{G18} No P{G19} No P{G20} No P{G21} No P{G31} No P{G32} No P{M-RlacZ} No P{tll(G1)lacZ} Yes P{tll(G2)lacZ} Yes P{tll(G3)lacZ} Yes P{tll(G4)lacZ} Yes P{tll(G5)lacZ} Yes P{tll(G6)lacZ} Yes P{tll-lacZ.K10} No P{tll-lacZ.K11} No P{tll-lacZ.K1} No P{tll-lacZ.K2} No P{tll-lacZ.K3} No P{tll-lacZ.K4} No P{tll-lacZ.K5} No P{tll-lacZ.K6} No P{tll-lacZ.K7} No P{tll-lacZ.K8} No P{tll-lacZ.K9} No P{tll-lacZ.P1} Yes P{tll-lacZ.P2} Yes P{tll-lacZ.P3} Yes P{tll-lacZ.P4} Yes P{tll-lacZ.S1} No P{tll-lacZ.S2} No P{tll-lacZ.S3} No P{tll-lacZ.S4} No P{tll-lacZ.S5} No P{tll-lacZ.S6} No P{tll-lacZ.S7} No P{tll-lacZ.Stat92mut} Yes UAS construct P{GD1548} NA P{UAS-tll.H} NA characterization construct P{D1.R} NA heat-shock construct P{HS-tll} NA P{hs-GAL4-tll.LBD.HA} NA P{tll-hsp70} NA
Insertions
	Type of insertions Name Expression data
Related Comments
	Please look at the allele reports for the complete phenotype data tll mutants display deletion of A8 and telson and a defective head skeleton. (Jurgens et al., 1984) A detailed analysis of the tll mutant phenotype has been carried out. (Strecker et al., 1986) Genetic and phenotypic investigation of the tll gene demonstrate it is required for the formation of the normal body pattern. tll and maternal terminal genes act together in a common pathway to establish the posterior and anterior domains of the embryo. (Strecker et al., 1988) Expression of kni-Ecol\lacZ regulatory fusion constructs in mutant embryos shows that kni expression is repressed by tll activity. (Pankratz et al., 1989) tll mutants exhibit deletions in the terminal regions of the embryo. (Gaul and Jaeckle, 1990) Molecular analysis and cloning of tll suggests that the tll protein functions as a transcription factor. (Pignoni et al., 1990) The effect of tll on hb and ftz expression has been studied. (Casanova, 1990) tll represses Kr expression in the central segmentation domain and activates Kr expression in the posterior Malpighian tubule domain. (Gaul and Weigel, 1990) tll was included in a study to determine how gap genes influence gt expression. (Eldon and Pirrotta, 1991) tll exerts a negative effect on gt expression in the embryo. (Kraut and Levine, 1991) Mutant tll embryos alter the expression of the Ubx bx region enhancer element, BRE. (Qian et al., 1991) Mutations in zygotic cardinal gene tll do not interact with RpII140wimp. (Parkhurst and Ish-Horowicz, 1991) The activation and spatial limitation of tll and hkb expression in the posterior region of the embryo is critically dependent on tor activity. The spatial limitation of hkb and tll expression is not regulated by the "central gap genes" which are essential for the establishment of segmentation in the trunk of the embryo, and also does not involve mutual interactions between hkb and tll. (Bronner and Jaeckle, 1991) Zygotically active locus involved in the terminal developmental program in the embryo. (Strecker et al., 1991) csw- syncytial and cellular blastoderm embryos show reduced posterior tll domain. csw and phl acts in concert to regulate tll expression. (Perkins et al., 1992) Dsor1 is involved in both the positive and negative regulation of tll. (Tsuda et al., 1993) An artificial bcd responder gene composed of three bcd consensus binding sites driving Ecol\lacZ is activated by bcd and repressed by tor. This repression does not require tll or hkb. (Ronchi et al., 1993) tll is required for normal pattern of prd in the embryo. (Gutjahr et al., 1993) Ecol\lacZ reporter gene constructs demonstrate the presence of tor, bcd and dl maternal system cis-acting response elements in the 5' flanking region of tll. (Liaw and Lengyel, 1993) "Splice" phenotype of torNRE dependent on tll, trk and tsl. (Casanova and Struhl, 1993) DNA sequence comparisons between D.melanogaster and D.virilis, DNAseI footprinting and promoter dissection identify two tll promoter elements in the previously defined D3 promoter region to be potential regulatory targets of phl-activated transcription factors. Sequence comparisons also reveal that unique residues in the DNA binding domain of the tll protein itself are conserved, and may be responsible for differences between the tll binding site and that of the closely related retinoid/estrogen receptors. (Liaw et al., 1993) The role of tll in the regulation of run mRNA expression in the early embryo has been investigated. (Klingler and Gergen, 1993) The BRE region of Ubx includes binding sites for hb, ftz, tll, en and twi. The binding of their products and the interplay between them is responsible for generating the expression pattern directed by the BRE. (Qian et al., 1993)