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General Information
Symbol
Dmel\tld
Species
D. melanogaster
Name
tolloid
Annotation Symbol
CG6868
Feature Type
FlyBase ID
FBgn0003719
Gene Model Status
Stock Availability
Gene Snapshot
tolloid (tld) encodes a member of the asticin metalloprotease family. Its primary function is to cleave the product of sog, which facilitates the diffusion of the ligand encoded by dpp to the dorsal-most cells in the early blastoderm embryo, helping to specify formation of the amnioserosa. [Date last reviewed: 2019-03-14]
Key Links
Genomic Location
Cytogenetic map
Sequence location
3R:24,749,731..24,753,510 [+]
Recombination map
3-85
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Molecular Function (GO)
[Detailed GO annotations]
Summaries
Pathway (FlyBase)
Positive Regulators of Activin Signaling Pathway -
Positive regulators of activin signaling up-regulate the pathway, ultimately resulting in the increased nuclear activity of the Smox/Med transcription factor complex.
Positive Regulators of BMP Signaling Pathway -
Positive regulators of Bone Morphogenetic Protein (BMP) signaling up-regulate the pathway, ultimately resulting in the increased nuclear activity of the Mad/Med transcription factor complex.
Protein Function (UniProtKB)
Required for normal dorsal development. TLD may interact physically with DPP-C protein.
(UniProt, P25723)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
tld: tolloid
Embryonic lethal. Partially ventralized. Cephalic furrow shifted dorsally; defects in germband extension. Denticle belts extended laterally.
Summary (Interactive Fly)
secreted BMP-1 homolog - member of the asticin metalloprotease family - cleaves Decapentaplegic inhibitor Short gastrulation thus functioning to activate Dpp - cleavage facilitates Dpp diffusion to the dorsal-most cells in the early blastoderm embryo, helping to specify formation of the amnioserosa
Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\tld or the JBrowse view of Dmel\tld for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Comments on Gene Model
Gene model reviewed during 5.49
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0084691
3359
1067
Additional Transcript Data and Comments
Reported size (kB)
3.5 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0084070
121.8
1067
6.94
Polypeptides with Identical Sequences

There is only one protein coding transcript and one polypeptide associated with this gene

Additional Polypeptide Data and Comments
Reported size (kDa)
1057 (aa); 116 (kD)
Comments
External Data
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\tld using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (13 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR001881, InterPro:IPR018097
(assigned by InterPro )
inferred from electronic annotation with InterPro:IPR006026
(assigned by InterPro )
Biological Process (8 terms)
Terms Based on Experimental Evidence (8 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
non-traceable author statement
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
inferred from direct assay
Terms Based on Predictions or Assertions (0 terms)
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
antennal anlage

Comment: reported as procephalic ectoderm anlage

central brain anlage

Comment: reported as procephalic ectoderm anlage

dorsal head epidermis anlage

Comment: reported as procephalic ectoderm anlage

visual anlage

Comment: reported as procephalic ectoderm anlage

antennal primordium

Comment: reported as procephalic ectoderm primordium

central brain primordium

Comment: reported as procephalic ectoderm primordium

visual primordium

Comment: reported as procephalic ectoderm primordium

dorsal head epidermis primordium

Comment: reported as procephalic ectoderm primordium

lateral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

ventral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

Additional Descriptive Data
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.
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.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Epitope-tagged tld protein is found in the dorsal half of the blastoderm embryo.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\tld in GBrowse 2
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 9-10
  • Stages(s) 11-12
  • Stages(s) 13-16
Alleles, Insertions, and Transgenic Constructs
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (10)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
10 of 15
Yes
Yes
 
10 of 15
Yes
Yes
8 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
 
2 of 15
No
No
1 of 15
No
No
1 of 15
Yes
No
1 of 15
Yes
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (10)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
11 of 15
Yes
Yes
11 of 15
Yes
Yes
10 of 15
Yes
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
Yes
No
Rattus norvegicus (Norway rat) (8)
11 of 13
Yes
Yes
9 of 13
No
Yes
5 of 13
No
Yes
2 of 13
No
No
2 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (15)
6 of 12
Yes
Yes
5 of 12
No
Yes
4 of 12
No
Yes
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
Yes
1 of 12
No
No
1 of 12
No
No
1 of 12
No
Yes
1 of 12
No
Yes
1 of 12
No
Yes
1 of 12
No
No
Danio rerio (Zebrafish) (18)
11 of 15
Yes
Yes
11 of 15
Yes
Yes
8 of 15
No
Yes
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (41)
11 of 15
Yes
Yes
3 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG0919016C )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila simulans
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091500OE )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Lucilia cuprina
Australian sheep blowfly
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W012C )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Rhodnius prolixus
Kissing bug
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Cimex lectularius
Bed bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X010N )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G009U )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (19)
5 of 10
4 of 10
3 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 1 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Interaction
    References
    Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
     
    Disease Associations of Human Orthologs (via DIOPT v7.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Functional Complementation Data
    Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
    Interactions
    Summary of Physical Interactions
    Summary of Genetic Interactions
    esyN Network Diagram
    esyN Network Key:
    Suppression
    Enhancement

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    External Data
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Gene Group - Pathway Membership (FlyBase)
    Positive Regulators of Activin Signaling Pathway -
    Positive regulators of activin signaling up-regulate the pathway, ultimately resulting in the increased nuclear activity of the Smox/Med transcription factor complex.
    Positive Regulators of BMP Signaling Pathway -
    Positive regulators of Bone Morphogenetic Protein (BMP) signaling up-regulate the pathway, ultimately resulting in the increased nuclear activity of the Mad/Med transcription factor complex.
    External Data
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    3R
    Recombination map
    3-85
    Cytogenetic map
    Sequence location
    3R:24,749,731..24,753,510 [+]
    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)
    96A-96A
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (9)
    Genomic Clones (14)
     

    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 sequences
    BDGP DGC clones
    Other clones
      Drosophila Genomics Resource Center cDNA clones

      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.

      cDNA Clones, End Sequenced (ESTs)
      BDGP DGC clones
      RNAi and Array Information
      Linkouts
      DRSC - Results frm RNAi screens
      GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
      Antibody Information
      Laboratory Generated Antibodies
       
      Commercially Available Antibodies
       
      Other Information
      Relationship to Other Genes
      Source for database identify of
      Source for identity of: tld CG6868
      Source for database merge of
      Additional comments
      Other Comments
      The tld product negatively regulates dac expression in the embryonic head.
      tld is required to keep the levels of sog protein low on the dorsal side of blastoderm embryos.
      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.
      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.
      Mutants show no interaction with Df(2R)Pcl11B or Df(3L)66C-G28.
      tld protein cleaves sog protein. Cleavage is stimulated by dpp protein.
      Mutations in spz, tld and yeti cooperatively induce brain tumor formation in the Malignant Brain Tumor strain.
      tld acts as a tumor suppressor.
      tld positively modulates dpp activity.
      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.
      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.
      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.
      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.
      tld alleles display relative phenotypic strengths, this may be correlated to the progressive loss of dorsal pattern elements in the ventralised mutants.
      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.
      The zygotically acting DV genes repress ac expression within specific DV domains.
      DNA sequence analysis of tld shows that the tkd gene product is closely related to the human bone marrow morphogenetic protein 1.
      tld is required for the normal ontogeny of the zen pattern and fating of the amnioserosa.
      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.
      Zygotically active locus involved in the terminal developmental program in the embryo.
      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.
      tld mutants display a twisted embryo, denticle belts are laterally spread.
      Origin and Etymology
      Discoverer
      Etymology
      Identification
      External Crossreferences and Linkouts ( 50 )
      Sequence Crossreferences
      NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
      GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
      GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
      RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
      UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
      UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
      Other crossreferences
      BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
      Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
      Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
      Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
      Flygut - An atlas of the Drosophila adult midgut
      GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
      iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
      KEGG Genes - Molecular building blocks of life in the genomic space.
      MEROPS - An information resource for peptidases (also termed proteases, proteinases and proteolytic enzymes) and the proteins that inhibit them.
      modMine - A data warehouse for the modENCODE project
      Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
      Linkouts
      BioGRID - A database of protein and genetic interactions.
      DroID - A comprehensive database of gene and protein interactions.
      DRSC - Results frm RNAi screens
      FLIGHT - Cell culture data for RNAi and other high-throughput technologies
      FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
      FlyMine - An integrated database for Drosophila genomics
      Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
      InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
      MIST (genetic) - An integrated Molecular Interaction Database
      MIST (protein-protein) - An integrated Molecular Interaction Database
      Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
      Synonyms and Secondary IDs (8)
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