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Gene Dmel\tok

General Information
Symbol	Dmel\tok	Species	D. melanogaster
Name	tolkin	Annotation symbol	CG6863
Feature type	protein_coding_gene	FlyBase ID	FBgn0004885
Gene Model Status	Current	Stock availability	11 publicly available
Also Known As	tlr, tlr-1, tldr-1
Genomic Location
Chromosome (arm)	3R	Recombination map	3-
Cytogenetic map	96A18-96A19	Sequence location	3R:20,545,419..20,574,855 [+]
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 tolkin is referred to in FlyBase by the symbol Dmel\tok (CG6863, FBgn0004885). 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: motor axon guidance; axon guidance; defasciculation of motor neuron axon; negative regulation of gene expression; imaginal disc-derived wing vein morphogenesis; protein processing. 36 alleles are reported. The phenotypes of these alleles are annotated with: organ system subdivision; adult segment; multi-cell-component structure; presumptive embryonic/larval nervous system; adult mesothoracic segment; peripheral nervous system; organ system; nervous system; hypodermal muscle of larval abdomen; abdominal segmental nerve. It has 2 annotated transcripts and 2 annotated polypeptides. 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 high expression to a trough of very low expression. Peak expression observed during late larval stages, during early pupal stages. Summary of FlyAtlas Anatomical Expression Data: Expression at high levels in the following post-embryonic organs or tissues: adult brain, larval/adult fat body, adult spermathecae. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, adult eye, larval/adult central nervous system, adult heart, larval/adult carcass. Comments on Affy2 ProbeSet: ProbeSet 1641053_s_at completely aligns to an exonic region common to each of the 2 FlyBase-annotated transcript isoforms of tok. Gene sequence location is 3R:20545419..20574855.
External Summaries
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	Sequence features BKN21716 HFA17087 Controlled Vocabulary Terms negative regulation of gene expression
FB2011_10	References Peluso et al., 2011, Dev. Cell 21(2): 375--383 Controlled Vocabulary Terms proteolysis protein processing
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 96A18-96A19   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 96A-96A     (O'Connor, 1994.7.26)
Experimentally Determined Recombination Data
Location	3- (O'Connor, 1994.7.26)
Left of (cM)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\tok 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) tok-RA FBtr0084690  NM_057531   5377   1464 tok-RB FBtr0084689  NM_170168   5255   1464
Additional Transcript Data & Comments
Reported size (kB)	5.5 (northern blot) (Finelli et al., 1995) 5.4 (northern blot) (Nguyen et al., 1994)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein tok-PA   FBpp0084069   165.2   1464   6.26     NP_476879   AAF56328 tok-PB   FBpp0084068   165.2   1464   6.26     NP_733035   AAN14017
Additional Polypeptide Data & Comments
Reported size (kDa)	1464 (aa); 165 (kD predicted) (Nguyen et al., 1994) 1464 (aa) (Finelli et al., 1995)
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 M12.011
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA698183   AE014297 AAF56328   AE014297 AAN14017   AC008205   AI257613   AI258193   AI293947   AI403694   AW940894   AY051585 AAK93009   AY061532 AAL29080   AY881252   AY881253   AY881254   AY881255 AAX98693   AY881256 AAX98694   AY881257 AAX98695   AY881258 AAX98696   AY881259 AAX98697   AY881260 AAX98698   AY881261 AAX98699   AY881262 AAX98700   AY881263   AY881264   AY881265   AY881266   AY881267   AY881268 AAX98701   AY881269 AAX98702   AY881270 AAX98703   AY881271   AY881272 AAX98704   AY881273 AAX98705   AY881274 AAX98706   AY881275 AAX98707   AY881276 AAX98708   AY881277 AAX98709   AY881278 AAX98710   AY881279 AAX98711   AY881280 AAX98712   AY881281 AAX98713   AY881282 AAX98714   AY881283 AAX98715   AY881284 AAX98716   AY881285 AAX98717   AY881286 AAX98718   AY886890   BF499804   BI571479   BI581919   BI582174   BI589553   BI591129   BI611827   BI621015   BI629960   CK131894   CK657291   CK660197   CL705868   CO188410   CO192928   CO266513   CO276717   CO279353   CO289524   CO293430   CO300649   CO323430   CO328690   CO329062   CZ471462   EC060185   EC062728   EC078262   EC086065   EC198459   EC198959   EC209341   EC215360   EC221631   EC228997   EC235699   EC242135   U12634 AAA70057   U34777 AAC47015
UniProtKB/Swiss-Prot
UniProtKB/TrEMBL	Q23995 Q24132 Q49II0 Q49II5 Q49II7 Q49II8 Q49IJ0 Q49IJ1 Q49IK4 Q95RA3 Q9VC47
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 10 -- 11   embryonic optic lobe primordium   (Nguyen et al., 1994) embryonic stage 14 -- 17   corpus allatum primordium   (Finelli et al., 1995, Nguyen et al., 1994) third instar larval stage   larval outer optic anlage   (Finelli et al., 1995) northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage   peripheral nervous system | restricted   (Nguyen et al., 1994) embryonic/larval pharynx   (Nguyen et al., 1994) amnioproctodeal invagination   (Nguyen et al., 1994) mesectoderm | vicinity of   (Finelli et al., 1995) stomodeal invagination   (Nguyen et al., 1994) embryonic stage -- adult stage     (Finelli et al., 1995, Nguyen et al., 1994) blastoderm stage   organism | dorsal   (Finelli et al., 1995) embryonic stage 5   organism | dorsal   (Nguyen et al., 1994) embryonic stage 10   peripheral nervous system precursor cluster   (Nguyen et al., 1994) embryonic stage 12 -- 17   somatic mesoderm   (Nguyen et al., 1994) visceral mesoderm   (Nguyen et al., 1994) embryonic stage 14 -- 17   central nervous system | restricted   (Finelli et al., 1995) embryonic stage 17   central nervous system | restricted   (Nguyen et al., 1994) sensillum basiconicum   (Nguyen et al., 1994) embryonic/larval proventriculus   (Nguyen et al., 1994) larval stage   embryonic/larval corpus allatum   (Nguyen et al., 1994) third instar larval stage   morphogenetic furrow | anterior to   (Finelli et al., 1995, Nguyen et al., 1994) eye-antennal disc   (Finelli et al., 1995, Nguyen et al., 1994) embryonic/larval brain   (Finelli et al., 1995) imaginal disc   (Finelli et al., 1995, Nguyen et al., 1994) embryonic/larval optic lobe   (Nguyen et al., 1994)
Additional Descriptive Data
	tok transcripts are expressed throughout development with the highest levels in larvae and pupae. By in situ hybridization, they are first detected on the dorsal surface of late cellular blastoderm embryos. In early stages of germ band extension, they are detected in the anterior and posterior midgut invaginations and later in the visceral mesoderm and pharynx. At stage 10, expression is observed in patches of thoracic cells thought to be proneural clusters and later strong expression is observed in thoracic PNS cells. At stage 10-11, transient expression is observed in optic lobe precursor cells. During dorsal closure, cells that will form the corpus allatum express tok. At stage 17, tok expression is observed in the CNS, in thoracic clusters thought to be the basiconical sensillum and in the proventriculus. In third instar larvae, tok is expressed in all imaginal discs and in the proliferative regions of the optic lobes. In the eye discs, strong tok expression is only found anterior to the morphogenetic furrow. Complex patterns of staining are seen in the other discs. Expression also continues in the corpus allatum of the ring gland. With the exception of the ring gland, the pattern of tok is very similar to that of tld in larval tissues but tld is expressed at a significantly lower level. (Nguyen et al., 1994) tok transcripts are expressed throughout development. They are first detected by in situ hybridization in the dorsal part of the blastoderm embryo. During germ band extension, tok is exressed near the mesectoderm. During stage 10, three faint lateral spots are observed in the thoracic segments. During dorsal closure, expression is seen in cells that will form the corpus allatum of the ring gland. Expression is also observed in a few cells in the CNS, a few of which were tentatively identified as RP neurons or just lateral to them. In later embryos, expression is observed in many large ventral cells in the nerve cord. In the larval brain, tok is expressed in distinct rings in both hemispheres, one of which corresponds to the outer proliferation center. tok stains most imaginal discs in their entirety with some areas staining more intensely than others. In the eye imaginal disc, the staining is located anterior to the morphogenetic furrow. A comparison of the expression patterns of tok, tld, and dpp is made. (Finelli et al., 1995)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
Additional Descriptive Data
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 FBgn0004885    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0004885 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 high expression to a trough of very low expression. Peak expression observed during late larval stages, during early pupal 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 FBgn0004885 expression level Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    755 embryo 04-06hr    608 embryo 06-08hr    1181 embryo 08-10hr    1543 embryo 10-12hr    1667 embryo 12-14hr    1907 embryo 14-16hr    1650 embryo 16-18hr    1059 embryo 18-20hr    1244 embryo 20-22hr    1204 embryo 22-24hr    1350 larva L1    1213 larva L2    622 larva L3 12hr old    1142 larva L3 puffstage 1-2    953 larva L3 puffstage 3-6    2194 larva L3 puffstage 7-9    4889 white prepupae new    7452 white prepupae 12hr    4918 white prepupae 24hr    2808 pupae 2d postWPP    2427 pupae 3d postWPP    1336 pupae 4d postWPP    3589 adult male 01day    1078 adult male 05day    873 adult male 30day    1004 adult female 01day    741 adult female 05day    542 adult female 30day    653 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr  (755) embryo 04-06hr  (608) embryo 06-08hr  (1181) embryo 08-10hr  (1543) embryo 10-12hr  (1667) embryo 12-14hr  (1907) embryo 14-16hr  (1650) embryo 16-18hr  (1059) embryo 18-20hr  (1244) embryo 20-22hr  (1204) embryo 22-24hr  (1350) larva L1  (1213) larva L2  (622) larva L3 12hr old  (1142) larva L3 puffstage 1-2  (953) larva L3 puffstage 3-6  (2194) larva L3 puffstage 7-9  (4889) white prepupae new  (7452) white prepupae 12hr  (4918) white prepupae 24hr  (2808) pupae 2d postWPP  (2427) pupae 3d postWPP  (1336) pupae 4d postWPP  (3589) adult male 01day  (1078) adult male 05day  (873) adult male 30day  (1004) adult female 01day  (741) adult female 05day  (542) adult female 30day  (653) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    755 embryo 04-06hr    608 embryo 06-08hr    1181 embryo 08-10hr  (1543) embryo 10-12hr  (1667) embryo 12-14hr  (1907) embryo 14-16hr  (1650) embryo 16-18hr    1059 embryo 18-20hr    1244 embryo 20-22hr    1204 embryo 22-24hr    1350 larva L1    1213 larva L2    622 larva L3 12hr old    1142 larva L3 puffstage 1-2    953 larva L3 puffstage 3-6  (2194) larva L3 puffstage 7-9  (4889) white prepupae new  (7452) white prepupae 12hr  (4918) white prepupae 24hr  (2808) pupae 2d postWPP  (2427) pupae 3d postWPP    1336 pupae 4d postWPP  (3589) adult male 01day    1078 adult male 05day    873 adult male 30day    1004 adult female 01day    741 adult female 05day    542 adult female 30day    653 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    755 embryo 04-06hr    608 embryo 06-08hr    1181 embryo 08-10hr    1543 embryo 10-12hr    1667 embryo 12-14hr    1907 embryo 14-16hr    1650 embryo 16-18hr    1059 embryo 18-20hr    1244 embryo 20-22hr    1204 embryo 22-24hr    1350 larva L1    1213 larva L2    622 larva L3 12hr old    1142 larva L3 puffstage 1-2    953 larva L3 puffstage 3-6    2194 larva L3 puffstage 7-9    4889 white prepupae new    7452 white prepupae 12hr    4918 white prepupae 24hr    2808 pupae 2d postWPP    2427 pupae 3d postWPP    1336 pupae 4d postWPP    3589 adult male 01day    1078 adult male 05day    873 adult male 30day    1004 adult female 01day    741 adult female 05day    542 adult female 30day    653 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    40 embryo 02-04hr    755 embryo 04-06hr    608 embryo 06-08hr    1181 embryo 08-10hr    1543 embryo 10-12hr    1667 embryo 12-14hr    1907 embryo 14-16hr    1650 embryo 16-18hr    1059 embryo 18-20hr    1244 embryo 20-22hr    1204 embryo 22-24hr    1350 larva L1    1213 larva L2    622 larva L3 12hr old    1142 larva L3 puffstage 1-2    953 larva L3 puffstage 3-6    2194 larva L3 puffstage 7-9    4889 white prepupae new    7452 white prepupae 12hr    4918 white prepupae 24hr    2808 pupae 2d postWPP    2427 pupae 3d postWPP    1336 pupae 4d postWPP    3589 adult male 01day    1078 adult male 05day    873 adult male 30day    1004 adult female 01day    741 adult female 05day    542 adult female 30day    653 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0004885 expression level Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    755 embryo 04-06hr    608 embryo 06-08hr    1181 embryo 08-10hr    1543 embryo 10-12hr    1667 embryo 12-14hr    1907 embryo 14-16hr    1650 embryo 16-18hr    1059 embryo 18-20hr    1244 embryo 20-22hr    1204 embryo 22-24hr    1350 larva L1    1213 larva L2    622 larva L3 12hr old    1142 larva L3 puffstage 1-2    953 larva L3 puffstage 3-6    2194 larva L3 puffstage 7-9    4889 white prepupae new    7452 white prepupae 12hr    4918 white prepupae 24hr    2808 pupae 2d postWPP    2427 pupae 3d postWPP    1336 pupae 4d postWPP    3589 adult male 01day    1078 adult male 05day    873 adult male 30day    1004 adult female 01day    741 adult female 05day    542 adult female 30day    653 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr  (755) embryo 04-06hr  (608) embryo 06-08hr  (1181) embryo 08-10hr  (1543) embryo 10-12hr  (1667) embryo 12-14hr  (1907) embryo 14-16hr  (1650) embryo 16-18hr  (1059) embryo 18-20hr  (1244) embryo 20-22hr  (1204) embryo 22-24hr  (1350) larva L1  (1213) larva L2  (622) larva L3 12hr old  (1142) larva L3 puffstage 1-2  (953) larva L3 puffstage 3-6  (2194) larva L3 puffstage 7-9  (4889) white prepupae new  (7452) white prepupae 12hr  (4918) white prepupae 24hr  (2808) pupae 2d postWPP  (2427) pupae 3d postWPP  (1336) pupae 4d postWPP  (3589) adult male 01day  (1078) adult male 05day  (873) adult male 30day  (1004) adult female 01day  (741) adult female 05day  (542) adult female 30day  (653) Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    755 embryo 04-06hr    608 embryo 06-08hr    1181 embryo 08-10hr  1543 embryo 10-12hr  1667 embryo 12-14hr  1907 embryo 14-16hr  1650 embryo 16-18hr    1059 embryo 18-20hr    1244 embryo 20-22hr    1204 embryo 22-24hr    1350 larva L1    1213 larva L2    622 larva L3 12hr old    1142 larva L3 puffstage 1-2    953 larva L3 puffstage 3-6  2194 larva L3 puffstage 7-9  (4889) white prepupae new  (7452) white prepupae 12hr  (4918) white prepupae 24hr  (2808) pupae 2d postWPP  2427 pupae 3d postWPP    1336 pupae 4d postWPP  (3589) adult male 01day    1078 adult male 05day    873 adult male 30day    1004 adult female 01day    741 adult female 05day    542 adult female 30day    653 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    755 embryo 04-06hr    608 embryo 06-08hr    1181 embryo 08-10hr    1543 embryo 10-12hr    1667 embryo 12-14hr    1907 embryo 14-16hr    1650 embryo 16-18hr    1059 embryo 18-20hr    1244 embryo 20-22hr    1204 embryo 22-24hr    1350 larva L1    1213 larva L2    622 larva L3 12hr old    1142 larva L3 puffstage 1-2    953 larva L3 puffstage 3-6    2194 larva L3 puffstage 7-9    4889 white prepupae new    7452 white prepupae 12hr    4918 white prepupae 24hr    2808 pupae 2d postWPP    2427 pupae 3d postWPP    1336 pupae 4d postWPP    3589 adult male 01day    1078 adult male 05day    873 adult male 30day    1004 adult female 01day    741 adult female 05day    542 adult female 30day    653 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    40 embryo 02-04hr    755 embryo 04-06hr    608 embryo 06-08hr    1181 embryo 08-10hr    1543 embryo 10-12hr    1667 embryo 12-14hr    1907 embryo 14-16hr    1650 embryo 16-18hr    1059 embryo 18-20hr    1244 embryo 20-22hr    1204 embryo 22-24hr    1350 larva L1    1213 larva L2    622 larva L3 12hr old    1142 larva L3 puffstage 1-2    953 larva L3 puffstage 3-6    2194 larva L3 puffstage 7-9    4889 white prepupae new    7452 white prepupae 12hr    4918 white prepupae 24hr    2808 pupae 2d postWPP    2427 pupae 3d postWPP    1336 pupae 4d postWPP    3589 adult male 01day    1078 adult male 05day    873 adult male 30day    1004 adult female 01day    741 adult female 05day    542 adult female 30day    653 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 FBgn0004885    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0004885 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Expression at high levels in the following post-embryonic organs or tissues: adult brain, larval/adult fat body, adult spermathecae. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, adult eye, larval/adult central nervous system, adult heart, larval/adult carcass. [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 FBgn0004885 expression level Tissue   Expression Level Larval Central Nervous System    353.225 Larval Midgut no informative data Larval Hindgut    45.8 Larval Malpighian Tubules    11.5 Larval Fat Body    615.4 Larval Salivary Gland    17.6 Larval Trachea no informative data Larval Carcass    307.3 Adult Head    397.8 Adult Eye    237.675 Adult Brain    716.8 Adult Thoracic-Abdominal Ganglion    257.7 Adult Crop no informative data Adult Midgut    7 Adult Hindgut    20.1 Adult Malpighian Tubules no informative data Adult Fat Body    922.6 Adult Salivary Gland    5.6 Adult Heart    369.25 Adult VirginFemale Spermatheca    791.3 Adult InseminatedFemale Spermatheca    614.5 Adult Ovary    14.5 Adult Testis    83.8 Adult Male Accessory Gland    4.6 Adult Carcass    284.5 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    353.225 Larval Midgut no informative data Larval Hindgut    45.8 Larval Malpighian Tubules    11.5 Larval Fat Body  (615.4) Larval Salivary Gland    17.6 Larval Trachea no informative data Larval Carcass    307.3 Adult Head    397.8 Adult Eye    237.675 Adult Brain  (716.8) Adult Thoracic-Abdominal Ganglion    257.7 Adult Crop no informative data Adult Midgut    7 Adult Hindgut    20.1 Adult Malpighian Tubules no informative data Adult Fat Body  (922.6) Adult Salivary Gland    5.6 Adult Heart    369.25 Adult VirginFemale Spermatheca  (791.3) Adult InseminatedFemale Spermatheca  (614.5) Adult Ovary    14.5 Adult Testis    83.8 Adult Male Accessory Gland    4.6 Adult Carcass    284.5 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    353.225 Larval Midgut no informative data Larval Hindgut    45.8 Larval Malpighian Tubules    11.5 Larval Fat Body    615.4 Larval Salivary Gland    17.6 Larval Trachea no informative data Larval Carcass    307.3 Adult Head    397.8 Adult Eye    237.675 Adult Brain    716.8 Adult Thoracic-Abdominal Ganglion    257.7 Adult Crop no informative data Adult Midgut    7 Adult Hindgut    20.1 Adult Malpighian Tubules no informative data Adult Fat Body    922.6 Adult Salivary Gland    5.6 Adult Heart    369.25 Adult VirginFemale Spermatheca    791.3 Adult InseminatedFemale Spermatheca    614.5 Adult Ovary    14.5 Adult Testis    83.8 Adult Male Accessory Gland    4.6 Adult Carcass    284.5 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    353.225 Larval Midgut no informative data Larval Hindgut    45.8 Larval Malpighian Tubules    11.5 Larval Fat Body    615.4 Larval Salivary Gland    17.6 Larval Trachea no informative data Larval Carcass    307.3 Adult Head    397.8 Adult Eye    237.675 Adult Brain    716.8 Adult Thoracic-Abdominal Ganglion    257.7 Adult Crop no informative data Adult Midgut    7 Adult Hindgut    20.1 Adult Malpighian Tubules no informative data Adult Fat Body    922.6 Adult Salivary Gland    5.6 Adult Heart    369.25 Adult VirginFemale Spermatheca    791.3 Adult InseminatedFemale Spermatheca    614.5 Adult Ovary    14.5 Adult Testis    83.8 Adult Male Accessory Gland    4.6 Adult Carcass    284.5 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to maximum FBgn0004885 expression level Tissue   Expression Level Larval Central Nervous System    353.225 Larval Midgut no informative data Larval Hindgut    45.8 Larval Malpighian Tubules    11.5 Larval Fat Body    615.4 Larval Salivary Gland    17.6 Larval Trachea no informative data Larval Carcass    307.3 Adult Head    397.8 Adult Eye    237.675 Adult Brain    716.8 Adult Thoracic-Abdominal Ganglion    257.7 Adult Crop no informative data Adult Midgut    7 Adult Hindgut    20.1 Adult Malpighian Tubules no informative data Adult Fat Body    922.6 Adult Salivary Gland    5.6 Adult Heart    369.25 Adult VirginFemale Spermatheca    791.3 Adult InseminatedFemale Spermatheca    614.5 Adult Ovary    14.5 Adult Testis    83.8 Adult Male Accessory Gland    4.6 Adult Carcass    284.5 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    353.225 Larval Midgut no informative data Larval Hindgut    45.8 Larval Malpighian Tubules    11.5 Larval Fat Body  615.4 Larval Salivary Gland    17.6 Larval Trachea no informative data Larval Carcass    307.3 Adult Head    397.8 Adult Eye    237.675 Adult Brain  716.8 Adult Thoracic-Abdominal Ganglion    257.7 Adult Crop no informative data Adult Midgut    7 Adult Hindgut    20.1 Adult Malpighian Tubules no informative data Adult Fat Body  (922.6) Adult Salivary Gland    5.6 Adult Heart    369.25 Adult VirginFemale Spermatheca  791.3 Adult InseminatedFemale Spermatheca  614.5 Adult Ovary    14.5 Adult Testis    83.8 Adult Male Accessory Gland    4.6 Adult Carcass    284.5 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    353.225 Larval Midgut no informative data Larval Hindgut    45.8 Larval Malpighian Tubules    11.5 Larval Fat Body    615.4 Larval Salivary Gland    17.6 Larval Trachea no informative data Larval Carcass    307.3 Adult Head    397.8 Adult Eye    237.675 Adult Brain    716.8 Adult Thoracic-Abdominal Ganglion    257.7 Adult Crop no informative data Adult Midgut    7 Adult Hindgut    20.1 Adult Malpighian Tubules no informative data Adult Fat Body    922.6 Adult Salivary Gland    5.6 Adult Heart    369.25 Adult VirginFemale Spermatheca    791.3 Adult InseminatedFemale Spermatheca    614.5 Adult Ovary    14.5 Adult Testis    83.8 Adult Male Accessory Gland    4.6 Adult Carcass    284.5 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    353.225 Larval Midgut no informative data Larval Hindgut    45.8 Larval Malpighian Tubules    11.5 Larval Fat Body    615.4 Larval Salivary Gland    17.6 Larval Trachea no informative data Larval Carcass    307.3 Adult Head    397.8 Adult Eye    237.675 Adult Brain    716.8 Adult Thoracic-Abdominal Ganglion    257.7 Adult Crop no informative data Adult Midgut    7 Adult Hindgut    20.1 Adult Malpighian Tubules no informative data Adult Fat Body    922.6 Adult Salivary Gland    5.6 Adult Heart    369.25 Adult VirginFemale Spermatheca    791.3 Adult InseminatedFemale Spermatheca    614.5 Adult Ovary    14.5 Adult Testis    83.8 Adult Male Accessory Gland    4.6 Adult Carcass    284.5 Expression Level Scale  None   Low   Moderate   High   Very high  Heatmap Tissue   Expression Level Larval Central Nervous System     Larval Midgut no informative data Larval Hindgut     Larval Malpighian Tubules     Larval Fat Body     Larval Salivary Gland     Larval Trachea no informative data Larval Carcass     Adult Head     Adult Eye     Adult Brain     Adult Thoracic-Abdominal Ganglion     Adult Crop no informative data Adult Midgut     Adult Hindgut     Adult Malpighian Tubules no informative data Adult Fat Body     Adult Salivary Gland     Adult Heart     Adult VirginFemale Spermatheca     Adult InseminatedFemale Spermatheca     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    397.8   NA  Eye    237.675   NA  Brain    716.8   353.225  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    257.7   NA  Crop    no informative data   no informative data  Midgut    7   45.8  Hindgut    20.1   11.5  Malpighian Tubules    no informative data   615.4  Fat Body    922.6   17.6  Salivary Gland    5.6   NA  Heart    369.25   no informative data  Trachea    NA   NA  VirginFemale Spermatheca    791.3   NA  InseminatedFemale Spermatheca    614.5   NA  Ovary    14.5   NA  Testis    83.8   NA  Male Accessory Gland    4.6   307.3  Carcass    284.5 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_25
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0004885 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG6863-RB FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for tok Stages(s) 4-6 Stages(s) 7-8 Stages(s) 13-16
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal tokΔ2-41 tokΔ3 tokΔ14 tokΔ24 tokΔ68 lethal | larval stage tokΔ2-41 lethal | larval stage (with Df(3R)tld-γ1) tokE1 lethal | larval stage (with tokE1) tokΔ2-41 lethal | larval stage (with tokΔ2-41) tokE1 lethal | larval stage | recessive tok2 tok3 tok4 tok5 tok6 tok7 lethal | pupal stage (with tokE1) tokΔ2-41 lethal | pupal stage (with tokΔ2-41) tokE1 viable tokEY00956 tokG3 viable, with Scer\GAL4pnr-MD237 tokGD245 Sterility Allele fertile tokEY00956 Other Phenotypes Allele developmental rate defective tokΔ2-41 neuroanatomy defective (with tokD427) tokK788 neuroanatomy defective (with tokK788) tokD427 neuroanatomy defective | embryonic stage tokΔ2-41 neuroanatomy defective | embryonic stage (with Df(3R)tld-γ1) tokE1 neuroanatomy defective | embryonic stage (with tokE1) tokΔ2-41 neuroanatomy defective | embryonic stage (with tokΔ2-41) tokE1 neuroanatomy defective | larval stage tokΔ2-41 neuroanatomy defective | larval stage (with Df(3R)tld-γ1) tokE1 neuroanatomy defective | larval stage (with tokE1) tokΔ2-41 neuroanatomy defective | larval stage (with tokΔ2-41) tokE1 visible, with Scer\GAL4how-24B tokScer\UAS.cSa.T:Ivir\HA1 Phenotype manifest in Allele A1-7 ventral longitudinal muscle 1 tokΔ2-41 A1-7 ventral longitudinal muscle 1 (with Df(3R)tld-γ1) tokE1 A1-7 ventral longitudinal muscle 1 (with tokE1) tokΔ2-41 A1-7 ventral longitudinal muscle 1 (with tokΔ2-41) tokE1 abdominal segmental nerve tokΔ2-41 abdominal segmental nerve (with Df(3R)tld-γ1) tokE1 abdominal segmental nerve (with tokE1) tokΔ2-41 abdominal segmental nerve (with tokΔ2-41) tokE1 anterior crossvein tokΔ2-41 crossvein (with tokE1) tokΔ2-41 crossvein (with tokΔ2-41) tokE1 embryonic/larval brain tok2 tok3 tok4 tok5 tok6 tok7 embryonic/larval neuromuscular junction tokΔ2-41 embryonic/larval neuromuscular junction (with Df(3R)tld-γ1) tokE1 embryonic/larval neuromuscular junction (with tokE1) tokΔ2-41 embryonic/larval neuromuscular junction (with tokΔ2-41) tokE1 imaginal disc | larval stage tok2 tok3 tok4 tok5 tok6 tok7 intersegmental nerve tokΔ2-41 intersegmental nerve (with Df(3R)tld-γ1) tokE1 intersegmental nerve (with tokD427) tokK788 intersegmental nerve (with tokE1) tokΔ2-41 intersegmental nerve (with tokK788) tokD427 intersegmental nerve (with tokΔ2-41) tokE1 intersegmental nerve | embryonic stage (with tokD427) tokK788 intersegmental nerve | embryonic stage (with tokK788) tokD427 larval neuromuscular junction & abdominal dorsal acute muscle 1 (with tokD427) tokK788 larval neuromuscular junction & abdominal dorsal acute muscle 1 (with tokK788) tokD427 larval neuromuscular junction & abdominal dorsal acute muscle 2 (with tokD427) tokK788 larval neuromuscular junction & abdominal dorsal acute muscle 2 (with tokK788) tokD427 larval neuromuscular junction & abdominal dorsal oblique muscle 1 (with tokD427) tokK788 larval neuromuscular junction & abdominal dorsal oblique muscle 1 (with tokK788) tokD427 larval neuromuscular junction & abdominal dorsal oblique muscle 2 (with tokD427) tokK788 larval neuromuscular junction & abdominal dorsal oblique muscle 2 (with tokK788) tokD427 larval neuromuscular junction & abdominal lateral transverse muscle 1 (with tokD427) tokK788 larval neuromuscular junction & abdominal lateral transverse muscle 1 (with tokK788) tokD427 larval neuromuscular junction & abdominal lateral transverse muscle 2 (with tokD427) tokK788 larval neuromuscular junction & abdominal lateral transverse muscle 2 (with tokK788) tokD427 larval neuromuscular junction & abdominal lateral transverse muscle 3 (with tokD427) tokK788 larval neuromuscular junction & abdominal lateral transverse muscle 3 (with tokK788) tokD427 larval neuromuscular junction & abdominal lateral transverse muscle 4 (with tokD427) tokK788 larval neuromuscular junction & abdominal lateral transverse muscle 4 (with tokK788) tokD427 larval neuromuscular junction & abdominal ventral acute muscle 1 (with tokD427) tokK788 larval neuromuscular junction & abdominal ventral acute muscle 1 (with tokK788) tokD427 larval neuromuscular junction & abdominal ventral acute muscle 2 (with tokD427) tokK788 larval neuromuscular junction & abdominal ventral acute muscle 2 (with tokK788) tokD427 larval neuromuscular junction & abdominal ventral acute muscle 3 (with tokD427) tokK788 larval neuromuscular junction & abdominal ventral acute muscle 3 (with tokK788) tokD427 larval neuromuscular junction & abdominal ventral longitudinal muscle 1 (with tokD427) tokK788 larval neuromuscular junction & abdominal ventral longitudinal muscle 1 (with tokK788) tokD427 larval neuromuscular junction & abdominal ventral longitudinal muscle 2 (with tokD427) tokK788 larval neuromuscular junction & abdominal ventral longitudinal muscle 2 (with tokK788) tokD427 larval neuromuscular junction & abdominal ventral longitudinal muscle 3 (with tokD427) tokK788 larval neuromuscular junction & abdominal ventral longitudinal muscle 3 (with tokK788) tokD427 larval neuromuscular junction & abdominal ventral longitudinal muscle 4 (with tokD427) tokK788 larval neuromuscular junction & abdominal ventral longitudinal muscle 4 (with tokK788) tokD427 larval neuromuscular junction & abdominal ventral oblique muscle 4 (with tokD427) tokK788 larval neuromuscular junction & abdominal ventral oblique muscle 4 (with tokK788) tokD427 larval neuromuscular junction & abdominal ventral oblique muscle 5 (with tokD427) tokK788 larval neuromuscular junction & abdominal ventral oblique muscle 5 (with tokK788) tokD427 larval neuromuscular junction & abdominal ventral oblique muscle 6 (with tokD427) tokK788 larval neuromuscular junction & abdominal ventral oblique muscle 6 (with tokK788) tokD427 longitudinal connective (with tokE1) tokΔ2-41 longitudinal connective (with tokΔ2-41) tokE1 posterior crossvein tokΔ2-41 presumptive embryonic/larval central nervous system (with tokD427) tokK788 presumptive embryonic/larval central nervous system (with tokK788) tokD427 segmental nerve (with tokD427) tokK788 segmental nerve (with tokK788) tokD427 segmental nerve | embryonic stage (with tokD427) tokK788 segmental nerve | embryonic stage (with tokK788) tokD427 wing, with Scer\GAL4how-24B tokScer\UAS.cSa.T:Ivir\HA1
Classical Alleles ( 26 )
For All Classical Alleles Show
Allele of tok Class Mutagen Stocks Known lesion tok1 amorphic allele - genetic evidence ethyl methanesulfonate 2 Yes tok3 PM hybrid dysgenesis 2 Yes tok5-HA-1069 1 -- tok5-HA-1856 1 -- tokd00218 P-element activity 1 -- tokEY00956 P-element activity 1 -- tokG3 P-element activity 1 Yes tokKG00406 1 -- tok2 PM hybrid dysgenesis 0 Yes tok4 PM hybrid dysgenesis 0 Yes tok5 PM hybrid dysgenesis 0 Yes tok6 PM hybrid dysgenesis 0 Yes tok7 PM hybrid dysgenesis 0 Yes tokD427 ethyl methanesulfonate 0 Yes tokE1 0 -- tokI678 ethyl methanesulfonate 0 Yes tokK598 ethyl methanesulfonate 0 Yes tokK788 ethyl methanesulfonate 0 Yes tokP natural population 0 Yes tokpir 0 -- tokunspecified 0 -- tokΔ14 Delta2-3 0 Yes tokΔ2-41 Delta2-3 0 Yes tokΔ24 Delta2-3 0 Yes tokΔ3 Delta2-3 0 Yes tokΔ68 Delta2-3 0 Yes
Alleles Carried on Transgenic Constructs ( 10 )
For All Alleles Carried on Transgenic Constructs Show
Allele of tok Class Mutagen Stocks Known lesion tokGD245 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes tokastTld.MtnA.T:Ivir\HA1 in vitro construct 0 Yes tokcd.MtnA.T:Ivir\HA1 in vitro construct - amino acid replacement 0 Yes tokcd.Scer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - amino acid replacement 0 Yes tokKK101665 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes tokMtnA.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes tokpm.MtnA.T:Ivir\HA1 in vitro construct - amino acid replacement 0 Yes tokpm.Scer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - amino acid replacement 0 Yes tokScer\UAS.cSa.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes toktld.PN in vitro construct - regulatory fusion 0 Yes
Aneuploid Aberrations
Not disrupted in	Df(3R)96B (Bloomington Drosophila Stock Center, 1998.6.2)
Disrupted in	Df(3R)BSC520 (Christensen et al., 2008.4.15) Df(3R)tld-γ1 (Finelli et al., 1995) Tp(3;2)slo3 (Nguyen et al., 1994)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD245} NA P{KK101665} NA P{UAS-tlr.cd} NA P{UAS-tlr.pm} NA P{UAS-tok.S} NA characterization construct P{tldP/tlr-1} NA
Insertions
	Type of insertions Name Expression data insertion of mobile activating element P{EPgy2}tokEY00956 NA P{XP}tokd00218 NA miscellaneous insertions P{RS5}tok5-HA-1069 NA P{RS5}tok5-HA-1856 NA P{SUPor-P}tokKG00406 NA P{wG3/4}tok2 NA P{wG3/4}tok3 NA P{wG3/4}tok4 NA P{wG3/4}tok5 NA P{wG3/4}tok6 NA P{wG3/4}tok7 NA P{wG3/4}tokG3 NA insertion of enhancer trap binary system P{GAL4}arl1 No
Gene Ontology: Function, Process & Cellular Component ( 10 unique terms )
Terms Based on Experimental Evidence ( 6 terms )
Molecular Function ( 0 terms)
Biological Process
CV term Evidence References
axon guidance inferred from mutant phenotype (Serpe and O'Connor, 2006) defasciculation of motor neuron axon inferred from mutant phenotype (Meyer and Aberle, 2006) imaginal disc-derived wing vein morphogenesis inferred from mutant phenotype (Serpe et al., 2005) motor axon guidance inferred from mutant phenotype (Meyer and Aberle, 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)
Cellular Component ( 0 terms)
Terms Based on Predictions or Assertions ( 4 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:P13497 (FlyBase, 1992-) zinc ion binding inferred from electronic annotation with InterPro:IPR006026 (FlyBase Curators et al., 2004-)
Biological Process
CV term Evidence References
proteolysis inferred from electronic annotation with InterPro:IPR001506, InterPro:IPR006026 (FlyBase Curators et al., 2004-)
Cellular Component ( 0 terms)
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 tok allele Gene References tokΔ2-41 sog (Araujo et al., 2003) unspecified daw (Serpe and O'Connor, 2006) sog (Serpe et al., 2005) tld (Serpe et al., 2005)
External Data
Linkouts	DPiM - DPiM, Drosophila Protein interaction Map • FBgn0004885 DroID - A comprehensive database of gene and protein interactions. • FBgn0004885 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 42944
Orthologs
Genome-wide drosophilid orthologs	Dana\GF17224
Curated drosophilid orthologs	Dpse\GA19912   Dsim\tok (Merritt et al., 2005.1.9)
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL013239-PA Aedes aegypti (mosquito) AGAP003702-PA Anopheles gambiae (mosquito) XP_393866.2 Apis mellifera (honey bee) 85253 Branchiostoma floridae (lancelet) ENSBTAP00000015593 Bos taurus (bovine) CN28118 Caenorhabditis brenneri (small nematode) CBP37016 Caenorhabditis briggsae (nematode) CE30977 Caenorhabditis elegans (worm) ENSCAFP00000013098 Canis familiaris (dog) ENSCINP00000007804 Ciona intestinalis (sea squirt) JA12969 Caenorhabditis japonica (nematode) CPIJ012475 Culex pipens (mosquito) ENSCPOP00000014968 Cavia porcellus (guinea pig) RP39340 Caenorhabditis remanei (nematode) ENSCSAVP00000017149 Ciona savignyi (sedentary tunicate) 221124 Capitella spl (polychaete worm) 204203 Daphnia pulex (daphnia) ENSDARP00000054471 Danio rerio (zebra fish) ENSECAP00000004160 Equus caballus (horse) ENSGACP00000022409 Gymnopilus aculeatus (mushroom) ENSGALP00000015567 Gallus gallus (chicken) 191792 Hypsipyla robusta (cedar shoot borer) ENSP00000061240 Homo sapiens (human) 236092 Lottia gigantea (owl limpet) ENSMODP00000002962 Monodelphis domestica (gray opossum) ENSMMUP00000034571 Macaca mulatta (rhesus monkey) ENSMUSP00000025986 Mus musculus (house mouse) 166027 Nematostella vectensis (starlet anemone) ENSOANP00000022827 Ornithorynchus anatinus (platypus) ENSORLP00000010047 Oryzias latipes (japanese killifish) PHUM003328-PA Pediculus humanus (human body louse) PP06600 Pristionchus pacificus (nematode) ENSPPYP00000002925 Pongo pygmaeus (orangutan) ENSPTRP00000028449 Pan troglodytes (chimpanzee) ENSRNOP00000018179 Rattus norvegicus (norway rat) Smp_134430 Schistosoma mansoni (trematode parasite) NP_999728.1 Strongylocentrotus purpuratus (purple sea urchin) XP_970162.1 Tribolium castaneum (red flour beetle) ENSTNIP00000009936 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000040418 Takifugu rubripes (pufferfish Takifugu) ENSXETP00000016336 Xenopus tropicalis (frog) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB10142 NV50061 LH21745 ISCW016869 FBgn0240688 FBgn0216039 FBgn0210748 FBgn0083414 FBgn0181474 JGI_V11_204203 FBgn0079908 FBgn0151226 FBgn0146745 FBgn0125913 FBgn0103614 FBgn0094242 CPIJ012475 BGIBMGA002519 ACYPI006588 GB10184 AGAP003702 AAEL013239 PHUM570110 GLEAN_11197
Stocks & Reagents
Stocks Listed in FlyBase ( 11 )
Bloomington	10922 P{wG3/4}tokG3, ry506 20105 y1 w67c23; P{EPgy2}tokEY00956 4569 cn1; tok3/TM3, Sb1 Ser1 4586 st1 e1 tok1/TM3, Sb1 Ser1 13657 y1 w67c23; ry506 P{SUPor-P}tokKG00406
Harvard	- P{XP}tokd00218
Kyoto	125047 w1118; P{RS5}tok5-HA-1069 125434 w1118; P{RS5}tok5-HA-1856 107808 st1 e1 tok1/TM3, Sb1 107800 cn1; tok3 / TM3, Sb1 Ser1
VDRC	v2656 w1118; P{GD245}v2656
Genomic Clones ( 2 )
	BACR33F18 BACR35N03 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 41 )
	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	GH23184 LP01328
Other clones	DPiM_GH23184
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	AT14557 HL03903 LP06026 LP07217 LP12952 LP18542 LP23474 RH02993 RH04849 RH06610 RH18250 RH19751 RH20126 RH52292 RH58907
Other clones	707 10828 19993 29876 33946 57701 63511 73451 161891 166636 CK01.8C.F3 CK01.90D.B10 EC34803 EK040028 EK071237 EK121911 EK134134 EK146640 EK178858 EK190444 EK202914 EK291210 EK291951
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0004885 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 42944
Antibody Information
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of	Source for identity of: tok CG6863 (Kodira, 1999.11)
Source for database merge of
Additional comments
Other Comments
	Located 700bp 5' to tld. (Finelli et al., 1995, Finelli et al., 1995) Point mutations and deletions in tok indicate it is vital during larval and pupal stages. Escapers of heteroallelic combinations have wing defects in which the posterior crossvein is missing. tok is expressed throughout embryogenesis in patterns that overlap and diverge from tld and dpp. (Finelli et al., 1995) tok is necessary for the proper development of the brain and imaginal discs and is involved in the specification of cell fates in the wing. (Finelli et al., 1995) The tld gene is the result of an ancient duplication that gave rise to the tld and tok genes, tok lies about 600bp 5' to tld. tok, like tld, is expressed throughout development and is mutatable to a lethal phenotype during larval stages. (Finelli et al., 1994) tok has been isolated, sequenced, expression pattern and mutant phenotype studied. tok is required for proper larval and pupal development. 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)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_057531 NM_170168 RefSeq (Proteins) • NP_476879 NP_733035 DDBJ / EMBL / GenBank • AA698183 AAA70057 AAC47015 AAF56328 AAK93009 AAL29080 AAN14017 AAX98693 AAX98694 AAX98695 AAX98696 AAX98697 AAX98698 AAX98699 AAX98700 AAX98701 AAX98702 AAX98703 AAX98704 AAX98705 AAX98706 AAX98707 AAX98708 AAX98709 AAX98710 AAX98711 AAX98712 AAX98713 AAX98714 AAX98715 AAX98716 AAX98717 AAX98718 AC008205 AI258193 AI403694 AW940894 AY051585 AY061532 AY881252 AY881253 AY881254 AY881255 AY881256 AY881257 AY881258 AY881259 AY881260 AY881261 AY881262 AY881263 AY881264 AY881265 AY881266 AY881267 AY881268 AY881269 AY881270 AY881271 AY881272 AY881273 AY881274 AY881275 AY881276 AY881277 AY881278 AY881279 AY881280 AY881281 AY881282 AY881283 AY881284 AY881285 AY881286 AY886890 CL705868 CZ471462 U12634 U34777 Entrez Gene - A searchable database of RefSeq genes. • 42944 UniProtKB/TrEMBL • Q9VC47 Q49II0 Q49II5 Q49II7 Q49II8 Q49IJ0 Q49IJ1 Q49IK4 Q95RA3 Q23995 Q24132
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 M12.011
Linkouts
	DPiM - DPiM, Drosophila Protein interaction Map • FBgn0004885 DroID - A comprehensive database of gene and protein interactions. • FBgn0004885 DRSC - Results from RNAi screens. • FBgn0004885 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0004885 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG6863-RB FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0004885 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 42944 InParanoid A subset of ortholog calls from InParanoid. • AAEL013239-PA Aedes aegypti (mosquito) AGAP003702-PA Anopheles gambiae (mosquito) XP_393866.2 Apis mellifera (honey bee) 85253 Branchiostoma floridae (lancelet) ENSBTAP00000015593 Bos taurus (bovine) CN28118 Caenorhabditis brenneri (small nematode) CBP37016 Caenorhabditis briggsae (nematode) CE30977 Caenorhabditis elegans (worm) ENSCAFP00000013098 Canis familiaris (dog) ENSCINP00000007804 Ciona intestinalis (sea squirt) JA12969 Caenorhabditis japonica (nematode) CPIJ012475 Culex pipens (mosquito) ENSCPOP00000014968 Cavia porcellus (guinea pig) RP39340 Caenorhabditis remanei (nematode) ENSCSAVP00000017149 Ciona savignyi (sedentary tunicate) 221124 Capitella spl (polychaete worm) 204203 Daphnia pulex (daphnia) ENSDARP00000054471 Danio rerio (zebra fish) ENSECAP00000004160 Equus caballus (horse) ENSGACP00000022409 Gymnopilus aculeatus (mushroom) ENSGALP00000015567 Gallus gallus (chicken) 191792 Hypsipyla robusta (cedar shoot borer) ENSP00000061240 Homo sapiens (human) 236092 Lottia gigantea (owl limpet) ENSMODP00000002962 Monodelphis domestica (gray opossum) ENSMMUP00000034571 Macaca mulatta (rhesus monkey) ENSMUSP00000025986 Mus musculus (house mouse) 166027 Nematostella vectensis (starlet anemone) ENSOANP00000022827 Ornithorynchus anatinus (platypus) ENSORLP00000010047 Oryzias latipes (japanese killifish) PHUM003328-PA Pediculus humanus (human body louse) PP06600 Pristionchus pacificus (nematode) ENSPPYP00000002925 Pongo pygmaeus (orangutan) ENSPTRP00000028449 Pan troglodytes (chimpanzee) ENSRNOP00000018179 Rattus norvegicus (norway rat) Smp_134430 Schistosoma mansoni (trematode parasite) NP_999728.1 Strongylocentrotus purpuratus (purple sea urchin) XP_970162.1 Tribolium castaneum (red flour beetle) ENSTNIP00000009936 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000040418 Takifugu rubripes (pufferfish Takifugu) ENSXETP00000016336 Xenopus tropicalis (frog) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /torstoll/tolodrel.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 42944 modMine - Data generated by the modENCODE project. • FBgn0004885 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB10142 NV50061 LH21745 ISCW016869 FBgn0240688 FBgn0216039 FBgn0210748 FBgn0083414 FBgn0181474 JGI_V11_204203 FBgn0079908 FBgn0151226 FBgn0146745 FBgn0125913 FBgn0103614 FBgn0094242 CPIJ012475 BGIBMGA002519 ACYPI006588 GB10184 AGAP003702 AAEL013239 PHUM570110 GLEAN_11197
Synonyms & Secondary IDs ( 19 )
Reported As
Symbol Synonym	CG6863 (Carrera et al., 2008, Keleman et al., 2009.8.5) CT21251 (Hynes and Zhao, 2000) tldr-1 (O'Connor, 1994.7.26, Padgett et al., 1992) tldr1   tlr (O'Connor et al., 2006, Serpe et al., 2005, Serpe and O'Connor, 2005, Araujo et al., 2003, Serpe and O'Connor, 2002, Serpe and O'Connor, 2006, Jensen et al., 2009, Peluso et al., 2011) Tlr (Whitman, 2005) tlr-1 (Aberle, 2005, Nguyen et al., 1994) tlr1 (Meyer and Aberle, 2006) Tlr-1 (Marques, 2005) Tok (Ashe, 2002) tok (Araujo et al., 2006, Carneiro et al., 2006, Anderson et al., 2006, Blanco et al., 2010, Christensen et al., 2008.4.15, Akdemir et al., 2007, Georlette et al., 2007, Liang et al., 2008, Carrera et al., 2008, Zeitlinger et al., 2007, Cook et al., 2010.2.12, Gim et al., 2001, Fritsch et al., 2010, Salzer et al., 2010) TOK (Banyai and Patthy, 2004)
Name Synonym	piranha (Meyer and Aberle, 2006) tld-related-1 (Nguyen et al., 1994) Tolkin (Araujo et al., 2005, Gim et al., 2001) tolkin (Finelli, 1995.8.23, Araujo et al., 2006, Carneiro et al., 2006, Anderson et al., 2006, Salzer et al., 2010) tolloid-related (Serpe and O'Connor, 2002, Serpe and O'Connor, 2006) Tolloid-related (Peluso et al., 2011) tolloid-related 1 (Padgett et al., 1992, Meyer and Aberle, 2006)
Secondary FlyBase IDs
References ( 65 )
Generate a list of	All references relating to this gene ( 65 )
List References by type	Research paper  ( 29 ) Supplementary material  ( 5 ) Review  ( 4 ) Personal communication to FlyBase  ( 6 ) Abstract  ( 8 ) FlyBase analysis  ( 3 ) Conference report  ( 1 ) DNA/RNA sequence record  ( 3 ) Curated genome annotation  ( 2 ) Computer file  ( 3 ) Automatic genome annotation  ( 1 )
Recent research papers ( 4 )
	Peluso et al., 2011, Dev. Cell 21(2): 375--383 Shaping BMP Morphogen Gradients through Enzyme-Substrate Interactions. [FBrf0214661] 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] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...