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Gene Dmel\tld

General Information
Symbol	Dmel\tld	Species	D. melanogaster
Name	tolloid	Annotation symbol	CG6868
Feature type	protein_coding_gene	FlyBase ID	FBgn0003719
Gene Model Status	Current	Stock availability	9 publicly available
Genomic Location
Chromosome (arm)	3R	Recombination map	3-85
Cytogenetic map	96A19-96A19	Sequence location	3R:20,575,455..20,579,232 [+]
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 tolloid is referred to in FlyBase by the symbol Dmel\tld (CG6868, FBgn0003719). It is a protein_coding_gene from Drosophila melanogaster. Based on sequence similarity, it is predicted to have molecular function: metalloendopeptidase activity. There is experimental evidence that it is involved in the biological process: torso signaling pathway; terminal region determination; negative regulation of gene expression; imaginal disc-derived wing vein morphogenesis; protein processing; maternal specification of dorsal/ventral axis, oocyte, soma encoded. 94 alleles are reported. The phenotypes of these alleles are annotated with: organ system; organ system subdivision; adult segment; portion of tissue; extended germ band embryo; adult mesothoracic segment; nervous system; non-connected developing system; organelle; primordium; epithelial furrow; external compound sense organ; late extended germ band embryo. It has one annotated transcript and one annotated polypeptide. Protein features are: Bone morphogenetic protein 1/tolloid-like protein; CUB; EGF-like calcium-binding; EGF-like calcium-binding, conserved site; EGF-like region, conserved site; EGF-type aspartate/asparagine hydroxylation site; Epidermal growth factor-like, type 3; Metallopeptidase, catalytic domain; Peptidase M12A, astacin; Peptidase, metallopeptidase. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of very low expression. Peak expression observed within 00-12 hour embryonic stages. Summary of FlyAtlas Anatomical Expression Data: Little or no expression detected in any larval or adult organs/tissues. Comments on Affy2 ProbeSet: ProbeSet 1624434_at completely aligns to an exonic region of the only FlyBase-annotated transcript isoform of tld. Gene sequence location is 3R:20575455..20579232.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	tld: tolloid Embryonic lethal. Partially ventralized. Cephalic furrow shifted dorsally; defects in germband extension. Denticle belts extended laterally.
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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FB2011_10	References Peluso et al., 2011, Dev. Cell 21(2): 375--383 Controlled Vocabulary Terms protein processing
FB2012_01	Sequence features BKN26256 HFA17084 Controlled Vocabulary Terms negative regulation of gene expression
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 96A19-96A19   Limits computationally determined from genome sequence between P{PZ}crb07207&P{PZ}BRWD305842 and P{EP}CycB3EP3127
Experimentally Determined Cytological Location
Cytogenetic map Notes References 96A21-96A21   (determined by in situ hybridisation)   (Ranz et al., 1999) 96A-96A   (determined by in situ hybridisation)   (Padgett and Gelbart, 1990) 96A21-96A21   (determined by in situ hybridisation)   (Shimell et al., 1991)
Experimentally Determined Recombination Data
Location	3-85 (Riede, 1996) 3-86 (Tearle and Nusslein-Volhard, 1987)
Left of (cM)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\tld for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) tld-RA FBtr0084691  NM_079763   3357   1067
Additional Transcript Data & Comments
Reported size (kB)	3.5 (northern blot) (Shimell et al., 1991)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein tld-PA   FBpp0084070   121.8   1067   6.94     NP_524487   AAF56329
Additional Polypeptide Data & Comments
Reported size (kDa)	1057 (aa); 116 (kD) (Shimell et al., 1991)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • EGF-like calcium-binding, conserved site (IPR018097) EGF-type aspartate/asparagine hydroxylation site (IPR000152) Epidermal growth factor-like, type 3 (IPR000742) CUB (IPR000859) Peptidase M12A, astacin (IPR001506) EGF-like calcium-binding (IPR001881) Peptidase, metallopeptidase (IPR006026) EGF-like region, conserved site (IPR013032) Bone morphogenetic protein 1/tolloid-like protein (IPR015446) Metallopeptidase, catalytic domain (IPR024079) MEROPS - An information resource for peptidases and the proteins that inhibit them • M12.010
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AE014297 AAF56329   AY886892   BI214737   BI228004   BI367501   BI368022   BI371067   BT010008 AAQ22477   CO297449   CO321362   EC051141   EC055663   EC200535   EC236403   GH756976   GH757385   GH759215   GH759225   GH760109   GH760676   GH773345   GH774017   GH775171   GH778088   GH781211   GH786538   GH786919   GH787816   GH789667   GH790458   GH790675   GH794957   GH801206   GH802775   GH802889   GH804489   GH810672   GH813057   GH814409   GH817683   GH818507   GH819731   GH819967   GH820002   GH822861   GH826774   GH828903   GH831110   GH833062   GH833992   GH835980   GH838258   GH840873   GH841312   GH843207   GH843369   GH843858   GH850236   GH851845   GH851847   GH855863   GH857800   GH859556   GH861909   GH863131   GH863336   GH863991   GH864551   GH866927   GH867384   GH868490   GH869385   GH870714   GH872162   GH873555   GH877410   GH877831   GH887711   GH887888   GH892106   GH894419   GH894760   GH895157   GH895225   GH895730   GH900097   GH910396   GH911417   GH915727   GH916136   GH916913   GH920173   GH922382   GH927068   GH927576   GH933412   GH934192   GH935259   GH940125   GH940690   GH940766   GH941064   M76976 AAA28491   U04239 AAC46482
UniProtKB/Swiss-Prot	P25723
UniProtKB/TrEMBL	Q7YU36
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
No Assay Recorded Stage Tissue/Position (including subcellular localization) Reference embryonic stage   tracheal pit   (Shimell et al., 1991) embryonic stage 8 -- 9   embryonic maxillary segment | restricted   (Shimell et al., 1991) embryonic stage 11   ventral neurogenic region | restricted   (Shimell et al., 1991) in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage | early   organism | dorsal   (Shimell et al., 1991) embryonic stage 4 -- 5   organism | dorsal & striped   (Zhang et al., 2001) embryonic stage 5     (Nguyen et al., 1994) embryonic stage 8 -- 9   embryonic labial segment | restricted   (Shimell et al., 1991) embryonic mandibular segment | restricted   (Shimell et al., 1991) third instar larval stage   imaginal disc   (Nguyen et al., 1994) morphogenetic furrow | anterior to   (Nguyen et al., 1994) embryonic/larval optic lobe   (Nguyen et al., 1994) eye-antennal disc   (Nguyen et al., 1994)
Additional Descriptive Data
	tld transcripts are observed in 0-12 hr embryos on northern blots. Transcripts are first detected by in situ hybridization at nuclear cycle 10-11 in dorsally located nuclei in the central portion of the embryo and in dorsally and ventrally positioned nuclei at the poles. The pattern extends and amplifies until at cycle 13, tld staining encompasses the dorsal-most 50% of the embryo. By the end of cellularization, four distinct bands of expression are observed. In late stage 8 to stage 9, a new pattern is seen. tld expression is observed in three lateral patches of cells in the emerging gnathal segments and more weakly in patches of cells in more posterior segments. At stage 11, the most intense staining is seen in a series of epidermal cells located ventral to each tracheal pit that are shaped like parentheses. Weaker staining is seen in cells encircling the tracheal pits and in patches of cells in the neurogenic ectoderm. (Shimell et al., 1991) tld expression is detected on the dorsal surface of cellular blastoderm embryos. It is detected in a similar pattern to that of tok but is detected at least 40 min. earlier. In third instar larvae, tld is expressed in a complex pattern in all imaginal discs and in the proliferative regions of the optic lobes. With the exception of its lack of expression in the ring gland, the pattern of tld is very similar to that of tok in larval tissues but tld is expressed at a significantly lower level. (Nguyen et al., 1994)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
No Assay Recorded Stage Tissue/Position (including subcellular localization) Reference blastoderm stage   organism | dorsal   (Marques et al., 1997)
Additional Descriptive Data
	Epitope-tagged tld protein is found in the dorsal half of the blastoderm embryo. (Marques et al., 1997)
Marker for
Subcellular Localization
CV Term
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 for FBgn0003719    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0003719 Very low expression bin max Moderate expression bin max High expression bin max Extremely high expression bin max Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of very low expression. Peak expression observed within 00-12 hour embryonic stages. [download data (TSV)] Guide to modENCODE expression level colors   No expression (0 - 0)   Extremely low expression (1 - 10)   Very low expression (11 - 100)   Low expression (101 - 400)   Moderate expression (401 - 1400)   Moderately high expression (1401 - 4000)   High expression (4001 - 10000)   Very high expression (10001 - 100000)   Extremely high expression (100001 - 2000000) Linear, scaled to maximum FBgn0003719 expression level Developmental Stage   Expression Level embryo 00-02hr    246 embryo 02-04hr    3429 embryo 04-06hr    1593 embryo 06-08hr    1836 embryo 08-10hr    1481 embryo 10-12hr    306 embryo 12-14hr    117 embryo 14-16hr    61 embryo 16-18hr    62 embryo 18-20hr    98 embryo 20-22hr    94 embryo 22-24hr    145 larva L1    77 larva L2    23 larva L3 12hr old    57 larva L3 puffstage 1-2    224 larva L3 puffstage 3-6    368 larva L3 puffstage 7-9    470 white prepupae new    348 white prepupae 12hr    619 white prepupae 24hr    294 pupae 2d postWPP    237 pupae 3d postWPP    116 pupae 4d postWPP    298 adult male 01day    46 adult male 05day    48 adult male 30day    69 adult female 01day    45 adult female 05day    41 adult female 30day    41 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (246) embryo 02-04hr  (3429) embryo 04-06hr  (1593) embryo 06-08hr  (1836) embryo 08-10hr  (1481) embryo 10-12hr  (306) embryo 12-14hr  (117) embryo 14-16hr    61 embryo 16-18hr    62 embryo 18-20hr    98 embryo 20-22hr    94 embryo 22-24hr  (145) larva L1    77 larva L2    23 larva L3 12hr old    57 larva L3 puffstage 1-2  (224) larva L3 puffstage 3-6  (368) larva L3 puffstage 7-9  (470) white prepupae new  (348) white prepupae 12hr  (619) white prepupae 24hr  (294) pupae 2d postWPP  (237) pupae 3d postWPP  (116) pupae 4d postWPP  (298) adult male 01day    46 adult male 05day    48 adult male 30day    69 adult female 01day    45 adult female 05day    41 adult female 30day    41 Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    246 embryo 02-04hr  (3429) embryo 04-06hr  (1593) embryo 06-08hr  (1836) embryo 08-10hr  1481 embryo 10-12hr    306 embryo 12-14hr    117 embryo 14-16hr    61 embryo 16-18hr    62 embryo 18-20hr    98 embryo 20-22hr    94 embryo 22-24hr    145 larva L1    77 larva L2    23 larva L3 12hr old    57 larva L3 puffstage 1-2    224 larva L3 puffstage 3-6    368 larva L3 puffstage 7-9    470 white prepupae new    348 white prepupae 12hr    619 white prepupae 24hr    294 pupae 2d postWPP    237 pupae 3d postWPP    116 pupae 4d postWPP    298 adult male 01day    46 adult male 05day    48 adult male 30day    69 adult female 01day    45 adult female 05day    41 adult female 30day    41 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    246 embryo 02-04hr    3429 embryo 04-06hr    1593 embryo 06-08hr    1836 embryo 08-10hr    1481 embryo 10-12hr    306 embryo 12-14hr    117 embryo 14-16hr    61 embryo 16-18hr    62 embryo 18-20hr    98 embryo 20-22hr    94 embryo 22-24hr    145 larva L1    77 larva L2    23 larva L3 12hr old    57 larva L3 puffstage 1-2    224 larva L3 puffstage 3-6    368 larva L3 puffstage 7-9    470 white prepupae new    348 white prepupae 12hr    619 white prepupae 24hr    294 pupae 2d postWPP    237 pupae 3d postWPP    116 pupae 4d postWPP    298 adult male 01day    46 adult male 05day    48 adult male 30day    69 adult female 01day    45 adult female 05day    41 adult female 30day    41 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    246 embryo 02-04hr    3429 embryo 04-06hr    1593 embryo 06-08hr    1836 embryo 08-10hr    1481 embryo 10-12hr    306 embryo 12-14hr    117 embryo 14-16hr    61 embryo 16-18hr    62 embryo 18-20hr    98 embryo 20-22hr    94 embryo 22-24hr    145 larva L1    77 larva L2    23 larva L3 12hr old    57 larva L3 puffstage 1-2    224 larva L3 puffstage 3-6    368 larva L3 puffstage 7-9    470 white prepupae new    348 white prepupae 12hr    619 white prepupae 24hr    294 pupae 2d postWPP    237 pupae 3d postWPP    116 pupae 4d postWPP    298 adult male 01day    46 adult male 05day    48 adult male 30day    69 adult female 01day    45 adult female 05day    41 adult female 30day    41 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0003719 expression level Developmental Stage   Expression Level embryo 00-02hr    246 embryo 02-04hr    3429 embryo 04-06hr    1593 embryo 06-08hr    1836 embryo 08-10hr    1481 embryo 10-12hr    306 embryo 12-14hr    117 embryo 14-16hr    61 embryo 16-18hr    62 embryo 18-20hr    98 embryo 20-22hr    94 embryo 22-24hr    145 larva L1    77 larva L2    23 larva L3 12hr old    57 larva L3 puffstage 1-2    224 larva L3 puffstage 3-6    368 larva L3 puffstage 7-9    470 white prepupae new    348 white prepupae 12hr    619 white prepupae 24hr    294 pupae 2d postWPP    237 pupae 3d postWPP    116 pupae 4d postWPP    298 adult male 01day    46 adult male 05day    48 adult male 30day    69 adult female 01day    45 adult female 05day    41 adult female 30day    41 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (246) embryo 02-04hr  (3429) embryo 04-06hr  (1593) embryo 06-08hr  (1836) embryo 08-10hr  (1481) embryo 10-12hr  (306) embryo 12-14hr  117 embryo 14-16hr    61 embryo 16-18hr    62 embryo 18-20hr    98 embryo 20-22hr    94 embryo 22-24hr  (145) larva L1    77 larva L2    23 larva L3 12hr old    57 larva L3 puffstage 1-2  (224) larva L3 puffstage 3-6  (368) larva L3 puffstage 7-9  (470) white prepupae new  (348) white prepupae 12hr  (619) white prepupae 24hr  (294) pupae 2d postWPP  (237) pupae 3d postWPP  116 pupae 4d postWPP  (298) adult male 01day    46 adult male 05day    48 adult male 30day    69 adult female 01day    45 adult female 05day    41 adult female 30day    41 Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    246 embryo 02-04hr  (3429) embryo 04-06hr  1593 embryo 06-08hr  1836 embryo 08-10hr  1481 embryo 10-12hr    306 embryo 12-14hr    117 embryo 14-16hr    61 embryo 16-18hr    62 embryo 18-20hr    98 embryo 20-22hr    94 embryo 22-24hr    145 larva L1    77 larva L2    23 larva L3 12hr old    57 larva L3 puffstage 1-2    224 larva L3 puffstage 3-6    368 larva L3 puffstage 7-9    470 white prepupae new    348 white prepupae 12hr    619 white prepupae 24hr    294 pupae 2d postWPP    237 pupae 3d postWPP    116 pupae 4d postWPP    298 adult male 01day    46 adult male 05day    48 adult male 30day    69 adult female 01day    45 adult female 05day    41 adult female 30day    41 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    246 embryo 02-04hr    3429 embryo 04-06hr    1593 embryo 06-08hr    1836 embryo 08-10hr    1481 embryo 10-12hr    306 embryo 12-14hr    117 embryo 14-16hr    61 embryo 16-18hr    62 embryo 18-20hr    98 embryo 20-22hr    94 embryo 22-24hr    145 larva L1    77 larva L2    23 larva L3 12hr old    57 larva L3 puffstage 1-2    224 larva L3 puffstage 3-6    368 larva L3 puffstage 7-9    470 white prepupae new    348 white prepupae 12hr    619 white prepupae 24hr    294 pupae 2d postWPP    237 pupae 3d postWPP    116 pupae 4d postWPP    298 adult male 01day    46 adult male 05day    48 adult male 30day    69 adult female 01day    45 adult female 05day    41 adult female 30day    41 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    246 embryo 02-04hr    3429 embryo 04-06hr    1593 embryo 06-08hr    1836 embryo 08-10hr    1481 embryo 10-12hr    306 embryo 12-14hr    117 embryo 14-16hr    61 embryo 16-18hr    62 embryo 18-20hr    98 embryo 20-22hr    94 embryo 22-24hr    145 larva L1    77 larva L2    23 larva L3 12hr old    57 larva L3 puffstage 1-2    224 larva L3 puffstage 3-6    368 larva L3 puffstage 7-9    470 white prepupae new    348 white prepupae 12hr    619 white prepupae 24hr    294 pupae 2d postWPP    237 pupae 3d postWPP    116 pupae 4d postWPP    298 adult male 01day    46 adult male 05day    48 adult male 30day    69 adult female 01day    45 adult female 05day    41 adult female 30day    41 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Developmental Stage   Expression Level embryo 00-02hr     embryo 02-04hr     embryo 04-06hr     embryo 06-08hr     embryo 08-10hr     embryo 10-12hr     embryo 12-14hr     embryo 14-16hr     embryo 16-18hr     embryo 18-20hr     embryo 20-22hr     embryo 22-24hr     larva L1     larva L2     larva L3 12hr old     larva L3 puffstage 1-2     larva L3 puffstage 3-6     larva L3 puffstage 7-9     white prepupae new     white prepupae 12hr     white prepupae 24hr     pupae 2d postWPP     pupae 3d postWPP     pupae 4d postWPP     adult male 01day     adult male 05day     adult male 30day     adult female 01day     adult female 05day     adult female 30day     FlyAtlas Anatomical Expression Data for FBgn0003719    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0003719 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Little or no expression detected in any larval or adult organs/tissues. [download data (TSV)] Guide to FlyAtlas expression level colors   No expression (0 - 9.999)   Low expression (10 - 99.999)   Moderate expression (100 - 499.999)   High level expression (500 - 999.999)   Very high expression (1000 - 25000) Linear, scaled to maximum FBgn0003719 expression level Tissue   Expression Level Larval Central Nervous System    48.475 Larval Midgut    1.8 Larval Hindgut    2.5 Larval Malpighian Tubules    2.2 Larval Fat Body no informative data Larval Salivary Gland    1.6 Larval Trachea    9.425 Larval Carcass no informative data Adult Head    9.8 Adult Eye    7.125 Adult Brain    22.4 Adult Thoracic-Abdominal Ganglion no informative data Adult Crop    1.5 Adult Midgut    1.9 Adult Hindgut    1.5 Adult Malpighian Tubules    4.4 Adult Fat Body no informative data Adult Salivary Gland    5.1 Adult Heart no informative data Adult VirginFemale Spermatheca    29.6 Adult InseminatedFemale Spermatheca no informative data Adult Ovary    1 Adult Testis    0.5 Adult Male Accessory Gland    1.4 Adult Carcass    2.7 Expression Level Scale  None   Low  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    48.475 Larval Midgut    1.8 Larval Hindgut    2.5 Larval Malpighian Tubules    2.2 Larval Fat Body no informative data Larval Salivary Gland    1.6 Larval Trachea    9.425 Larval Carcass no informative data Adult Head    9.8 Adult Eye    7.125 Adult Brain    22.4 Adult Thoracic-Abdominal Ganglion no informative data Adult Crop    1.5 Adult Midgut    1.9 Adult Hindgut    1.5 Adult Malpighian Tubules    4.4 Adult Fat Body no informative data Adult Salivary Gland    5.1 Adult Heart no informative data Adult VirginFemale Spermatheca    29.6 Adult InseminatedFemale Spermatheca no informative data Adult Ovary    1 Adult Testis    0.5 Adult Male Accessory Gland    1.4 Adult Carcass    2.7 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    48.475 Larval Midgut    1.8 Larval Hindgut    2.5 Larval Malpighian Tubules    2.2 Larval Fat Body no informative data Larval Salivary Gland    1.6 Larval Trachea    9.425 Larval Carcass no informative data Adult Head    9.8 Adult Eye    7.125 Adult Brain    22.4 Adult Thoracic-Abdominal Ganglion no informative data Adult Crop    1.5 Adult Midgut    1.9 Adult Hindgut    1.5 Adult Malpighian Tubules    4.4 Adult Fat Body no informative data Adult Salivary Gland    5.1 Adult Heart no informative data Adult VirginFemale Spermatheca    29.6 Adult InseminatedFemale Spermatheca no informative data Adult Ovary    1 Adult Testis    0.5 Adult Male Accessory Gland    1.4 Adult Carcass    2.7 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    48.475 Larval Midgut    1.8 Larval Hindgut    2.5 Larval Malpighian Tubules    2.2 Larval Fat Body no informative data Larval Salivary Gland    1.6 Larval Trachea    9.425 Larval Carcass no informative data Adult Head    9.8 Adult Eye    7.125 Adult Brain    22.4 Adult Thoracic-Abdominal Ganglion no informative data Adult Crop    1.5 Adult Midgut    1.9 Adult Hindgut    1.5 Adult Malpighian Tubules    4.4 Adult Fat Body no informative data Adult Salivary Gland    5.1 Adult Heart no informative data Adult VirginFemale Spermatheca    29.6 Adult InseminatedFemale Spermatheca no informative data Adult Ovary    1 Adult Testis    0.5 Adult Male Accessory Gland    1.4 Adult Carcass    2.7 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to maximum FBgn0003719 expression level Tissue   Expression Level Larval Central Nervous System    48.475 Larval Midgut    1.8 Larval Hindgut    2.5 Larval Malpighian Tubules    2.2 Larval Fat Body no informative data Larval Salivary Gland    1.6 Larval Trachea    9.425 Larval Carcass no informative data Adult Head    9.8 Adult Eye    7.125 Adult Brain    22.4 Adult Thoracic-Abdominal Ganglion no informative data Adult Crop    1.5 Adult Midgut    1.9 Adult Hindgut    1.5 Adult Malpighian Tubules    4.4 Adult Fat Body no informative data Adult Salivary Gland    5.1 Adult Heart no informative data Adult VirginFemale Spermatheca    29.6 Adult InseminatedFemale Spermatheca no informative data Adult Ovary    1 Adult Testis    0.5 Adult Male Accessory Gland    1.4 Adult Carcass    2.7 Expression Level Scale  None   Low  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    48.475 Larval Midgut    1.8 Larval Hindgut    2.5 Larval Malpighian Tubules    2.2 Larval Fat Body no informative data Larval Salivary Gland    1.6 Larval Trachea    9.425 Larval Carcass no informative data Adult Head    9.8 Adult Eye    7.125 Adult Brain    22.4 Adult Thoracic-Abdominal Ganglion no informative data Adult Crop    1.5 Adult Midgut    1.9 Adult Hindgut    1.5 Adult Malpighian Tubules    4.4 Adult Fat Body no informative data Adult Salivary Gland    5.1 Adult Heart no informative data Adult VirginFemale Spermatheca    29.6 Adult InseminatedFemale Spermatheca no informative data Adult Ovary    1 Adult Testis    0.5 Adult Male Accessory Gland    1.4 Adult Carcass    2.7 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    48.475 Larval Midgut    1.8 Larval Hindgut    2.5 Larval Malpighian Tubules    2.2 Larval Fat Body no informative data Larval Salivary Gland    1.6 Larval Trachea    9.425 Larval Carcass no informative data Adult Head    9.8 Adult Eye    7.125 Adult Brain    22.4 Adult Thoracic-Abdominal Ganglion no informative data Adult Crop    1.5 Adult Midgut    1.9 Adult Hindgut    1.5 Adult Malpighian Tubules    4.4 Adult Fat Body no informative data Adult Salivary Gland    5.1 Adult Heart no informative data Adult VirginFemale Spermatheca    29.6 Adult InseminatedFemale Spermatheca no informative data Adult Ovary    1 Adult Testis    0.5 Adult Male Accessory Gland    1.4 Adult Carcass    2.7 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    48.475 Larval Midgut    1.8 Larval Hindgut    2.5 Larval Malpighian Tubules    2.2 Larval Fat Body no informative data Larval Salivary Gland    1.6 Larval Trachea    9.425 Larval Carcass no informative data Adult Head    9.8 Adult Eye    7.125 Adult Brain    22.4 Adult Thoracic-Abdominal Ganglion no informative data Adult Crop    1.5 Adult Midgut    1.9 Adult Hindgut    1.5 Adult Malpighian Tubules    4.4 Adult Fat Body no informative data Adult Salivary Gland    5.1 Adult Heart no informative data Adult VirginFemale Spermatheca    29.6 Adult InseminatedFemale Spermatheca no informative data Adult Ovary    1 Adult Testis    0.5 Adult Male Accessory Gland    1.4 Adult Carcass    2.7 Expression Level Scale  None   Low   Moderate   High   Very high  Heatmap Tissue   Expression Level Larval Central Nervous System     Larval Midgut     Larval Hindgut     Larval Malpighian Tubules     Larval Fat Body no informative data Larval Salivary Gland     Larval Trachea     Larval Carcass no informative data Adult Head     Adult Eye     Adult Brain     Adult Thoracic-Abdominal Ganglion no informative data Adult Crop     Adult Midgut     Adult Hindgut     Adult Malpighian Tubules     Adult Fat Body no informative data Adult Salivary Gland     Adult Heart no informative data Adult VirginFemale Spermatheca     Adult InseminatedFemale Spermatheca no informative data Adult Ovary     Adult Testis     Adult Male Accessory Gland     Adult Carcass     FlyAtlas Organ/Tissue Expression, larval vs. adult Larval Expression Level Tissue Adult Expression Level   NA  Head    9.8   NA  Eye    7.125   NA  Brain    22.4   48.475  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    no informative data   NA  Crop    1.5   1.8  Midgut    1.9   2.5  Hindgut    1.5   2.2  Malpighian Tubules    4.4   no informative data  Fat Body    no informative data   1.6  Salivary Gland    5.1   NA  Heart    no informative data   9.425  Trachea    NA   NA  VirginFemale Spermatheca    29.6   NA  InseminatedFemale Spermatheca    no informative data   NA  Ovary    1   NA  Testis    0.5   NA  Male Accessory Gland    1.4   no informative data  Carcass    2.7 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_31
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003719 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG6868-RA FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for tld Stages(s) 4-6 Stages(s) 7-8 Stages(s) 9-10 Stages(s) 11-12 Stages(s) 13-16
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal (with tld1) tld14 lethal (with tld14) tld1 lethal | embryonic stage | heat sensitive tld10 tld12 lethal | embryonic stage | recessive tld1 tld2 tld3 tld5 tld6 tld7 tld12 tld14 tldB1 tldB2 tldB3 tldB4 tldB5 tldB6 tldB7 tldB8 tldE4 tldE5 tldE6 tldE7 tldE8 tldE9 tldE10 tldE11 tldE14 tldE15 tldunspecified lethal | heat sensitive tld10 lethal | pupal stage | temperature conditional (with tld1) tldMBT lethal | pupal stage | temperature conditional (with tldMBT) tld1 lethal | recessive tld18 tld19 tld20 tld21 tld22 tld23 viable, with Scer\GAL4pnr-MD237 tldGD246 Other Phenotypes Allele visible, with Scer\GAL4A9 tldA.Scer\UAS.T:Ivir\HA1 visible (with tld1) tldMBT visible (with tldMBT) tld1 wild-type tld+mNa tld+t12 tld+t19 Phenotype manifest in Allele amnioserosa tldB6 brain (with tld1) tldMBT brain (with tldMBT) tld1 cephalic furrow tldunspecified denticle belt tldunspecified denticle belt | maternal effect | somatic clone tldunspecified dorsal appendage | maternal effect | somatic clone tldunspecified egg operculum | maternal effect | somatic clone tldunspecified embryo | dorsal tldunspecified embryonic/larval dorsal vessel tld14 embryonic dorsal epidermis | maternal effect | somatic clone tldunspecified embryonic epidermis tld1 tld12 tld14 tld2 tld3 tld5 tld6 tld7 tldE10 tldE11 tldE14 tldE15 tldE4 tldE6 tldE7 tldE8 tldE9 embryonic leading edge cell tld9 embryonic ventral epidermis | maternal effect | somatic clone tldunspecified epidermis | dorsal tldB6 extended germ band embryo tldunspecified ommatidium (with tld1) tldMBT ommatidium (with tldMBT) tld1 polytene chromosome tldMBT posterior crossvein, with Scer\GAL4A9 tldScer\UAS.cHa posterior crossvein, with Scer\GAL4da.G32 tldScer\UAS.cHa posterior crossvein, with Scer\GAL4en-e16E tldA.Scer\UAS.T:Ivir\HA1 tldScer\UAS.cHa visceral mesoderm tld14 wing, with Scer\GAL4A9 tldA.Scer\UAS.T:Ivir\HA1 wing vein, with Scer\GAL4A9 tldA.Scer\UAS.T:Ivir\HA1
Classical Alleles ( 63 )
For All Classical Alleles Show
Allele of tld Class Mutagen Stocks Known lesion tld17 ethyl methanesulfonate 2 -- tldB4 amorphic allele - genetic evidence 1 Yes tld14 antimorphic allele - genetic evidence, loss of function allele ethyl methanesulfonate 1 Yes tld2 loss of function allele ethyl methanesulfonate 1 Yes tld9 ethyl methanesulfonate 1 -- tld15 amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes tld4 amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes tld5 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes tld68 amorphic allele - genetic evidence 0 -- tld6P117 amorphic allele - genetic evidence 0 Yes tldB6 amorphic allele - genetic evidence 0 -- tld10 ethyl methanesulfonate 0 Yes tld10E5.1A 0 Yes tld10E5.6I 0 Yes tld10E5.7E 0 Yes tld10E5.7G 0 Yes tld10E5H10B X ray 0 Yes tld11 ethyl methanesulfonate 0 -- tld12 antimorphic allele - genetic evidence ethyl methanesulfonate 0 Yes tld13 ethyl methanesulfonate 0 -- tld16 ethyl methanesulfonate 0 -- tld18 ethyl methanesulfonate 0 -- tld19 ethyl methanesulfonate 0 -- tld1 antimorphic allele - genetic evidence ethyl methanesulfonate 0 Yes tld20 ethyl methanesulfonate 0 -- tld21 ethyl methanesulfonate 0 -- tld22 ethyl methanesulfonate 0 -- tld23 ethyl methanesulfonate 0 -- tld3 antimorphic allele - genetic evidence ethyl methanesulfonate 0 Yes tld502 0 -- tld6 antimorphic allele - genetic evidence ethyl methanesulfonate 0 Yes tld6P11 0 -- tld6P1 0 -- tld6P41 antimorphic allele - genetic evidence 0 Yes tld7074 0 -- tld7 loss of function allele ethyl methanesulfonate 0 Yes tld7O47 0 -- tld8 ethyl methanesulfonate 0 Yes tld9K88 0 -- tld9Q19 antimorphic allele - genetic evidence 0 Yes tld9Q74 0 -- tldB1 0 Yes tldB2 0 Yes tldB3 antimorphic allele - genetic evidence 0 Yes tldB5 0 -- tldB7 0 Yes tldB8 0 -- tldE10 loss of function allele ethyl methanesulfonate 0 Yes tldE11 antimorphic allele - genetic evidence ethyl methanesulfonate 0 Yes tldE14 loss of function allele ethyl methanesulfonate 0 Yes tldE15 antimorphic allele - genetic evidence ethyl methanesulfonate 0 Yes tldE1 ethyl methanesulfonate 0 Yes tldE4 loss of function allele ethyl methanesulfonate 0 Yes tldE5 ethyl methanesulfonate 0 Yes tldE6 loss of function allele ethyl methanesulfonate 0 Yes tldE7 loss of function allele ethyl methanesulfonate 0 Yes tldE8 loss of function allele ethyl methanesulfonate 0 Yes tldE9 loss of function allele ethyl methanesulfonate 0 Yes tldMBT 0 -- tldP natural population 0 Yes tldr1 0 -- tldr2 0 -- tldunspecified ethyl methanesulfonate 0 --
Alleles Carried on Transgenic Constructs ( 31 )
For All Alleles Carried on Transgenic Constructs Show
Allele of tld Class Mutagen Stocks Known lesion tldGD246 in vitro construct - RNAi in vitro construct - regulatory fusion 2 Yes tldKK106256 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes tld+mNa in vitro construct - minigene 0 Yes tld+t12 in vitro construct - genomic fragment 0 Yes tld+t19 in vitro construct - genomic fragment 0 Yes tldA.MtnA.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - site directed mutagenesis in vitro construct - deletion 0 Yes tldA.MtnA.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - site directed mutagenesis in vitro construct - deletion 0 Yes tldA.Scer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldAct5C.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldAct5C.T:SV5\V5,T:Zzzz\His6 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldAo.MtnA.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - site directed mutagenesis in vitro construct - deletion 0 Yes tldAo.MtnA.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - site directed mutagenesis in vitro construct - deletion 0 Yes tldastTlr.MtnA.T:Ivir\HA1 in vitro construct 0 Yes tldD581A.Act5C.T:SV5\V5 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldD581E.Act5C.T:SV5\V5 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldE835K.Act5C.T:Ivir\HA1 in vitro construct - amino acid replacement 0 Yes tldMtnA.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldMtnA.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldN760I.Act5C.T:Ivir\HA1 in vitro construct - amino acid replacement 0 Yes tldo.MtnA.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - site directed mutagenesis 0 Yes tldo.MtnA.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - site directed mutagenesis 0 Yes tldpm.MtnA.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - site directed mutagenesis 0 Yes tldS915L.Act5C.T:Ivir\HA1 in vitro construct - amino acid replacement 0 Yes tldScer\UAS.cHa in vitro construct - regulatory fusion 0 Yes tldScer\UAS.cUa 0 Yes tldScer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldtld.T:Ivir\HA1 in vitro construct - coding region fusion 0 Yes tldtrunc.Act5C.T:SV5\V5,T:Zzzz\His6 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldΔEGF1+2.Act5C.T:SV5\V5,T:Zzzz\His6 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldΔEGF1.Act5C.T:SV5\V5,T:Zzzz\His6 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tldΔEGF2.Act5C.T:SV5\V5,T:Zzzz\His6 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(3R)96B (Bloomington Drosophila Stock Center, 1998.6.2) Df(3R)B204 (Jurgens et al., 1984) Df(3R)B219 (Muller et al., 1999) Df(3R)BSC397 (Christensen et al., 2007.10.29) Df(3R)BSC492 (Christensen et al., 2008.4.15) Df(3R)BSC519 (Christensen et al., 2008.4.15) Df(3R)BSC520 (Christensen et al., 2008.4.15) Df(3R)ED6085 (Ryder and Roote, 2004.11.10) Df(3R)ED6220 (Ryder and Roote, 2004.11.10) Df(3R)Hdγ1 (Saunders et al., 1997) Df(3R)tld68-62 (Shimell et al., 1991, Childs and O'Connor, 1994, Nguyen et al., 1994) Df(3R)tld-γ1 (Finelli et al., 1994, Finelli et al., 1995, Adamson and Shearn, 1996) Df(3R)X18E (Riede, 2001) Df(3R)XM54 (Jurgens et al., 1984) Df(3R)XTA1 (Jacobs et al., 1998, Riede, 2001, Tearle and Nusslein-Volhard, 1987, Ferguson and Anderson, 1992) Tp(3;2)slo3 (Nguyen et al., 1994) Ts(YLt;3Lt)G73 (Jurgens et al., 1984) Ts(YSt;3Lt)H173 (Jurgens et al., 1984)
Not disrupted in	Df(3R)ash18 (Adamson and Shearn, 1996) Df(3R)crb87-5 (Adamson and Shearn, 1996)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD246} NA P{KK106256} NA P{UAS-tld.A.T:HA1} NA P{UAS-tld.H} NA P{UAS-tld.T:HA1} NA P{UAS-tld.U} NA reporter construct P{tld423m-MSE-eve-lacZ} No P{tld423-MSE-eve-lacZ} No P{tld645m-MSE-eve-lacZ} No P{tld645-MSE-eve-lacZ} No P{tld-lacZ} No P{tolloid-lacZ.eve.-42.mut} No P{tolloid-lacZ.eve.-42} No characterization construct P{tld+12} NA P{tld+19} NA P{tldtld.T:HA1} NA P{tldP/tld} NA
Insertions
	Type of insertions Name Expression data
Gene Ontology: Function, Process & Cellular Component ( 16 unique terms )
Terms Based on Experimental Evidence ( 6 terms )
Molecular Function ( 0 terms)
Biological Process
CV term Evidence References
imaginal disc-derived wing vein morphogenesis inferred from mutant phenotype (Serpe et al., 2005) maternal specification of dorsal/ventral axis, oocyte, soma encoded inferred from mutant phenotype (Carneiro et al., 2006) negative regulation of gene expression inferred from direct assay (Anderson et al., 2006) protein processing inferred from direct assay (Serpe and O'Connor, 2006) terminal region determination inferred from genetic interaction with tor (Strecker et al., 1991) torso signaling pathway inferred from genetic interaction with tor (Strecker et al., 1991)
Cellular Component ( 0 terms)
Terms Based on Predictions or Assertions ( 10 terms )
Molecular Function
CV term Evidence References
calcium ion binding inferred from electronic annotation with InterPro:IPR001881, InterPro:IPR018097 (FlyBase Curators et al., 2004-) metalloendopeptidase activity inferred from sequence or structural similarity with UniProtKB:P98063 (FlyBase, 1992-) non-traceable author statement (Ray and Wharton, 2001, Lall and Patel, 2001) zinc ion binding inferred from electronic annotation with InterPro:IPR006026 (FlyBase Curators et al., 2004-)
Biological Process
CV term Evidence References
amnioserosa formation non-traceable author statement (Stronach and Perrimon, 2001) embryonic pattern specification traceable author statement (Chang and Hemmati-Brivanlou, 1998) positive regulation of BMP signaling pathway traceable author statement (Ray and Wharton, 2001) proteolysis inferred from electronic annotation with InterPro:IPR001506, InterPro:IPR006026 (FlyBase Curators et al., 2004-) non-traceable author statement (Chang and Hemmati-Brivanlou, 1998) regulation of transforming growth factor beta receptor signaling pathway traceable author statement (Lall and Patel, 2001) zygotic specification of dorsal/ventral axis non-traceable author statement (Swiss-Prot Project Members, 1992.5.1)
Cellular Component
CV term Evidence References
extracellular region non-traceable author statement (Chang and Hemmati-Brivanlou, 1998)
Sequence Ontology: Class of 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 tld allele Gene References tld1 sog (Ferguson and Anderson, 1992) tld14 dpp (Raftery et al., 1995) sog (Ferguson and Anderson, 1992) tldA.Scer\UAS.T:Ivir\HA1 tsg (Shimmi and O'Connor, 2003) tldE1 dpp (Raftery et al., 1995) tldMBT spz (Riede, 1996) tldunspecified sog (Ferguson and Anderson, 1992) unspecified dpp (Finelli et al., 1994, Finelli et al., 1995) sog (Shimmi and O'Connor, 2003, Serpe et al., 2005) tok (Serpe et al., 2005) yeti (Riede, 1996)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 67870 DroID - A comprehensive database of gene and protein interactions. • FBgn0003719 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 42945
Orthologs
Genome-wide drosophilid orthologs	Dana\GF23044
Curated drosophilid orthologs	Dbuz\tld (Ranz et al., 1999) Dmau\tld (Tao et al., 2003, Tao et al., 2003) Dpse\GA19917   Drep\tld (Ranz et al., 1999) Dsim\tld (Begun, 2000.4.5) Dvir\tld (Ranz et al., 1999)
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • XP_001943348.1 Acyrthosiphon pisum (pea aphid) BGIBMGA002519-PA Bombyx mori (silk worm) ISCW016869-PA Ixodes scapularis (deer tick) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • FBgn0240689 FBgn0216040 FBgn0210749 FBgn0041655 FBgn0248222 FBgn0151227 FBgn0146746 FBgn0125914 FBgn0103615 FBgn0100038
Stocks & Reagents
Stocks Listed in FlyBase ( 9 )
Bloomington	35502 st1 e1 tld2/TM3, Sb1 34041 st1 e1 tld14/TM3, Sb1 3303 Pr1 Dr1/TM3, Sb1 tld17 Ser1 5396 tldB4 pp/TM3, P{Thb8-lacZ}WD1, Sb1 Ser1 3237 ru1 h1 th1 st1 cu1 sr1 es tld9 ca1/TM3, Sb1 Ser1
Kyoto	106968 Pr1 Dr1/TM3, Sb1 tld17
VDRC	v100930 P{KK106256}VIE-260B v1215 w1118; P{GD246}v1215 v1216 w1118; P{GD246}v1216
Genomic Clones ( 1 )
	BACR33F18 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 99 )
	Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see GBrowse for alignment of the cDNAs and ESTs to the gene model.
cDNA Clones, Fully Sequenced
BDGP DGC clones	RE25412
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	RE20851 RE53275 RE53923 RE57880
Other clones	61982 83255 EC051141 EC055663 EK185526 EK217014 RP003023085 RP003086115 RP003090856 RP003101115 RP003106559 RP003117204 RP003129983 RP003150144 RP003151473 RP003157751 RP003168232 RP003174631 RP003177293 RP003179381 RP003180716 RP003184058 RP003188252 RP003198978 RP003203728 RP003222020 RP003223874 RP003224010 RP003227552 RP003228926 RP003230627 RP003239413 RP003242206 RP003244743 RP003246933 RP003251027 RP003255080 RP003261738 RP003274427 RP003283766 RP003284320 RP003294295 RP003294837 RP003308411 RP003311604 RP003314188 RP003321882 RP003325038 RP003325048 RP003325985 RP003334830 RP003345535 RP003355195 RP003359461 RP003361200 RP003384434 RP003406135 RP003408061 RP003420995 RP003424172 RP003448297 RP003452857 RP003458269 RP003471988 RP003486040 RP003487166 RP003489408 RP003500574 RP003502008 RP003508668 RP003515624 RP003530216 RP003532950 RP003538086 RP003540604 RP003557208 RP003566388 RP003582184 RP003590007 RP003626283 RP003628575 RP003633656 RP003635858 RP003638719 RP003638961 RP003644101 RP003644241 RP003656736 RP003661974 RP003662642 RP003668582 RP003669955 RP003670201 RP003679442
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0003719 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 42945
Antibody Information
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of	Source for identity of: tld CG6868 (Kodira, 1999.11)
Source for database merge of
Additional comments
Other Comments
	The tld product negatively regulates dac expression in the embryonic head. (Anderson et al., 2006) tld is required to keep the levels of sog protein low on the dorsal side of blastoderm embryos. (Srinivasan et al., 2002) In a sample of 79 genes with multiple introns, 33 showed significant heterogeneity in G+C content among introns of the same gene and significant positive correspondence between the intron and the third codon position G+C content within genes. These results are consistent with selection adding against preferred codons at the start of genes. (Kliman and Eyre-Walker, 1998) Mutants are isolated in an EMS mutagenesis screen to identify zygotic mutations affecting germ cell migration at discrete points during embryogenesis: mutants exhibit dorsal/ventral polarity pattern defects. (Moore et al., 1998) Mutants show no interaction with Df(2R)Pcl11B or Df(3L)66C-G28. (Nicholls and Gelbart, 1998) tld protein cleaves sog protein. Cleavage is stimulated by dpp protein. (Marques et al., 1997) Mutations in spz, tld and yeti cooperatively induce brain tumor formation in the Malignant Brain Tumor strain. (Riede, 1996) tld acts as a tumor suppressor. (Riede, 1996) tld positively modulates dpp activity. (Arora et al., 1995) Genetic and phenotypic data reveals that tld is involved in post-transcriptional regulation of dpp activity by elevating dpp activity dorsally. Mutations in tld disrupt amnioserosa formation. Injection of dpp RNA into the ventral side results in an asymmetric response, lateral expansion of the preexisting dorsal amnioserosa rather than formation of a second amnioserosa at the site of injection. (Neul and Ferguson, 1995) The tld protein appears to function by forming a complex containing dpp product, via protein-interacting EGF and C1r/s domains. The protease activity of the tld product is necessary, either directly or indirectly, for the activation of the dpp complex. (Finelli et al., 1994) 800bp of promoter sequences are sufficient for the spatial regulation of tld, determined using Ecol\lacZ reporter gene constructs. dl directly represses tld gene expression in ventral regions of the early embryo. DNaseI footprinting reveals dl binds to at least three sites in the tld 5' flanking sequences that are required in vivo for ventral repression (423bp VRE element). These sequences can behave as a transcriptional silencer to confer repression on a heterologous promoter. (Kirov et al., 1994) tok and tld arose by an ancient gene duplication event and each has subsequently evolved stage specific functions so they are no longer functionally equivalent. (Nguyen et al., 1994) tld alleles display relative phenotypic strengths, this may be correlated to the progressive loss of dorsal pattern elements in the ventralised mutants. (Arora and Nusslein-Volhard, 1992) tld, srw and sog are required to generate a gradient of dpp activity which directly specifies the pattern of the dorsal 40% of the embryo. Wild type tld is required for normal patterning only during the cellularisation of the blastoderm. (Ferguson and Anderson, 1992) The zygotically acting DV genes repress ac expression within specific DV domains. (Skeath et al., 1992) DNA sequence analysis of tld shows that the tkd gene product is closely related to the human bone marrow morphogenetic protein 1. (Childs et al., 1991) tld is required for the normal ontogeny of the zen pattern and fating of the amnioserosa. (Ray et al., 1991) Mutant analysis reveals that tld is required for normal dorsal development. Sequence analysis of tld suggests significant conservation of molecular mechanisms between mammals and insects. (Shimell et al., 1991) Zygotically active locus involved in the terminal developmental program in the embryo. (Strecker et al., 1991) Involved in the regulatory hierarchy responsible for the asymmetric distribution and function of zygotic regulatory gene products along the DV axis of early embryos. tld is required for the maintenance and refinement of the zen expression pattern during cellularization and gastrulation. (Rushlow and Levine, 1990) tld mutants display a twisted embryo, denticle belts are laterally spread. (Jurgens et al., 1984)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_079763 RefSeq (Proteins) • NP_524487 DDBJ / EMBL / GenBank • AAA28491 AAC46482 AAF56329 AAQ22477 AY886892 BI228004 BT010008 M76976 U04239 Entrez Gene - A searchable database of RefSeq genes. • 42945 UniProtKB/Swiss-Prot • P25723 UniProtKB/TrEMBL • Q7YU36
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • EGF-like calcium-binding, conserved site (IPR018097) EGF-type aspartate/asparagine hydroxylation site (IPR000152) Epidermal growth factor-like, type 3 (IPR000742) CUB (IPR000859) Peptidase M12A, astacin (IPR001506) EGF-like calcium-binding (IPR001881) Peptidase, metallopeptidase (IPR006026) EGF-like region, conserved site (IPR013032) Bone morphogenetic protein 1/tolloid-like protein (IPR015446) Metallopeptidase, catalytic domain (IPR024079) MEROPS - An information resource for peptidases and the proteins that inhibit them • M12.010
Linkouts
	BioGRID - A database of protein and genetic interactions • 67870 DroID - A comprehensive database of gene and protein interactions. • FBgn0003719 DRSC - Results from RNAi screens. • FBgn0003719 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003719 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG6868-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0003719 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 42945 InParanoid A subset of ortholog calls from InParanoid. • XP_001943348.1 Acyrthosiphon pisum (pea aphid) BGIBMGA002519-PA Bombyx mori (silk worm) ISCW016869-PA Ixodes scapularis (deer tick) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /torstoll/tolloid.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 42945 modMine - Data generated by the modENCODE project. • FBgn0003719 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • FBgn0240689 FBgn0216040 FBgn0210749 FBgn0041655 FBgn0248222 FBgn0151227 FBgn0146746 FBgn0125914 FBgn0103615 FBgn0100038 REDfly - A database of transcriptional regulatory elements. • FBgn0003719
Synonyms & Secondary IDs ( 8 )
Reported As
Symbol Synonym	CG6868 (Gonzalez-Aguero et al., 2005, Keleman et al., 2009.8.5) CT21278 (Hynes and Zhao, 2000) dTld (Lee et al., 2009) Tld (Wang and Ferguson, 2004, Ray and Wharton, 2001, Dale, 2000, Mizutani et al., 2005, Marques, 2005, Umulis et al., 2010, Peluso et al., 2011) tld (Araujo et al., 2006, Umulis et al., 2006, Carneiro et al., 2006, Zeitlinger et al., 2007, Canty et al., 2006, Anderson et al., 2006, Stathopoulos and Levine, 2005, Christensen et al., 2007.10.29, Serpe and O'Connor, 2006, Blanco et al., 2010, Meyer and Aberle, 2006, Christensen et al., 2008.4.15, Christensen et al., 2008.4.15, Christensen et al., 2008.4.15, Lilja et al., 2007, Liang et al., 2008, Fritsch et al., 2010, Salzer et al., 2010, Pruteanu-Malinici et al., 2011) TLD (Banyai and Patthy, 2004) Tolloid (Araujo et al., 2005)
Name Synonym	tolloid (Araujo et al., 2006, Carneiro et al., 2006, Canty et al., 2006, Anderson et al., 2006, Lilja et al., 2007, Salzer et al., 2010, Zhang et al., 2001) Tolloid (Banyai and Patthy, 2004, Lee et al., 2009, Umulis et al., 2010, Peluso et al., 2011)
Secondary FlyBase IDs
References ( 198 )
Generate a list of	All references relating to this gene ( 198 )
List References by type	Research paper  ( 91 ) Supplementary material  ( 5 ) Review  ( 53 ) Personal communication to FlyBase  ( 10 ) Abstract  ( 27 ) FlyBase analysis  ( 1 ) Stock list  ( 1 ) Curated genome annotation  ( 2 ) Protein sequence record  ( 1 ) Computer file  ( 2 ) DNA/RNA sequence record  ( 1 ) Automatic genome annotation  ( 1 ) Book  ( 1 ) Conference report  ( 1 )
Recent research papers ( 6 )
	Peluso et al., 2011, Dev. Cell 21(2): 375--383 Shaping BMP Morphogen Gradients through Enzyme-Substrate Interactions. [FBrf0214661] Pruteanu-Malinici et al., 2011, PLoS Comput. Biol. 7(7): e1002098 Automatic Annotation of Spatial Expression Patterns via Sparse Bayesian Factor Models. [FBrf0214618] Blanco et al., 2010, BMC Dev. Biol. 10: 94 Gene expression following induction of regeneration in Drosophila wing imaginal discs. Expression profile of regenerating wing discs. [FBrf0211812] Fritsch et al., 2010, Dev. Genes Evol. 220(9-10): 235--250 Rapid evolution of a novel signalling mechanism by concerted duplication and divergence of a BMP ligand and its extracellular modulators. [FBrf0212674] Salzer et al., 2010, Genetics 184(1): 185--197 The retinal determination gene eyes absent is regulated by the EGF receptor pathway throughout development in Drosophila. [FBrf0209690] Umulis et al., 2010, Dev. Cell 18(2): 260--274 Organism-Scale Modeling of Early Drosophila Patterning via Bone Morphogenetic Proteins. [FBrf0210054] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...