tlr, tolloid-related 1, tolloid-related, tlr-1, tldr-1
BMP-1 homolog - a Zinc metallopeptidase with similar substrate specificity to tolloid, which is its paralog. Tok cleaves the Decapentaplegic inhibitor Short gastrulation as well as the prodomains of several TGF-beta ligands
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.49
Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\tok using the Feature Mapper tool.
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.
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.
GBrowse - Visual display of RNA-Seq signalsView Dmel\tok in GBrowse 2
Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
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.
For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.
Source for identity of: tok CG6863
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.
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.
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.
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.