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General Information
Symbol
Dmel\gd
Species
D. melanogaster
Name
gastrulation-defective
Annotation Symbol
CG1505
Feature Type
FlyBase ID
FBgn0000808
Gene Model Status
Stock Availability
Gene Snapshot
gastrulation-defective (gd) encodes a secreted serine protease. It is a member of the protease cascade that generates the ligand that activates the receptor encoded by Tl. It contributes to establishment of the dorsal-ventral embryonic axis. [Date last reviewed: 2019-03-07]
Also Known As
fs(1)M18, SP186, fs(1)573, fs(1)gd
Key Links
Genomic Location
Cytogenetic map
Sequence location
X:11,985,154..11,987,764 [-]
Recombination map
1-38
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
Belongs to the peptidase S1 family. (O62589)
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
Predictions / Assertions
Summaries
Gene Group (FlyBase)
DORSAL GROUP -
The Dorsal group genes encode components of the Toll pathway and were initially identified by maternal effect mutations resulting in the dorsalization of the embryo. (Adapted from FBrf0091014 and FBrf0223077).
S1A SERINE PROTEASES - CHYMOTRYPSIN-LIKE -
Chymotrypsin-like serine proteases cleave substrates following one of the hydrophobic amino acids at the P1 position. (Adapted from FBrf0239469). Chymotrypsin-like serine proteases of the S1A family are endopeptidases that cleave substrates following one of the hydrophobic amino acids at the P1 position. (Adapted from FBrf0239469).
Pathway (FlyBase)
Extracellular Spatzle Activating Pathway Core Components -
In Drosophila, activation of the Toll pathway is controlled by the generation of a cleaved, active, Toll-binding form of spatzle ligand. Proteolytic activation of spatzle ligand lies downstream of several zymogen activation cascades that are initiated by different cues. In dorsal-ventral patterning, localized activation of spz results in ventral nuclear accumulation of dl. During viral and fungal and gram positive bacterial immune challenge, zymogen cascades are activated by extracellular pattern recognition receptors or virulence factor-mediated cleavage of the zymogen persephone (psh). (Adapted from FBrf0091014 and FBrf0223077).
Protein Function (UniProtKB)
Component of the extracellular signaling pathway that establishes the dorsal-ventral pathway of the embryo (PubMed:9618496, PubMed:9477324, PubMed:12493753). Three proteases; ndl, gd and snk process easter to create active easter (PubMed:9477324). Active easter defines cell identities along the dorsal-ventral continuum by activating the spz ligand for the Tl receptor in the ventral region of the embryo (PubMed:9477324).
(UniProt, O62589)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
gd: gastrulation defective
Maternal-effect lethal; embryos produced by homozygous females exhibit hyperplasia of dorsal cuticular elements and aplasia of ventral elements; phenotye of strong alleles, e.g. gd7, indistinguishable from that of dl. Weaker alleles display some ventral elements. Gastrula exhibits excessive furrowing both dorsally and ventrally [Zalokar, Audit, and Erk, 1975, Dev. Biol. 47: 419-32 (fig.)]. Polarity of the egg shell unaffected. Temperature-sensitive period of temperature-sensitive alleles begins several hours prior to ovoposition and persists until 1.5 h after fertilization. Five of eight mutants classed as weak on the basis of filzkorper development; these five tend to be more dorsalized in posterior than in anterior parts of the embryo, as the setal bands become progressively narrowed posteriorly. gd7/gd7; Toll/+ females produce lateralized embryos; most ventral and dorsal pattern elements missing; lateral cephalic furrow seen both dorsally and ventrally, and ventral setae formed at all dorsal-ventral positions.
Summary (Interactive Fly)
a maternally supplied protease involved in establishment of dorsal/ventral polarity - targets Snake
Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
3

Please see the GBrowse view of Dmel\gd or the JBrowse view of Dmel\gd 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 5.51
Gene model reviewed during 5.45
Gene model reviewed during 5.41
Gene model reviewed during 5.39
Gene model reviewed during 6.03
Alternative translation stop created by use of multiphasic reading frames within coding region.
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0332308
1914
528
FBtr0332309
1923
531
FBtr0332310
1932
179
FBtr0347416
2234
528
Additional Transcript Data and Comments
Reported size (kB)
2.1 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0304587
59.1
528
7.31
FBpp0304588
59.5
531
7.31
FBpp0304589
20.5
179
9.91
FBpp0312566
59.1
528
7.31
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

528 aa isoforms: gd-PC, gd-PF
Additional Polypeptide Data and Comments
Reported size (kDa)
528 (aa); 34, 30, 28 (kD observed); 59 (kD predicted)
Comments
External Data
Crossreferences
InterPro - A database of protein families, domains and functional sites
MEROPS - An information resource for peptidases (also termed proteases, proteinases and proteolytic enzymes) and the proteins that inhibit them.
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\gd 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 (8 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from sequence model
inferred from sequence or structural similarity with HGNC:3551
Biological Process (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
inferred from genetic interaction with FLYBASE:ea; FB:FBgn0000533
inferred from genetic interaction with FLYBASE:pip; FB:FBgn0003089
inferred from mutant phenotype
(assigned by UniProt )
inferred from genetic interaction with FLYBASE:pip; FB:FBgn0003089
inferred from genetic interaction with FLYBASE:Tl; FB:FBgn0262473
inferred from direct assay
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:Spn27A; FB:FBgn0028990
inferred from direct assay
inferred from mutant phenotype
(assigned by UniProt )
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
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
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
gd transcripts are observed exclusively in ovaries and early embryos on northern blots. gd transcripts are first observed in the germarium in nurse cells. As oogenesis proceeds they accumulate in nurse cells and in the oocyte.By stage 10 they are observed in follicle cells as well.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
gd protein is detected in ovaries and early embryos on western blots. Levels are highest in ovaries and it is essentially gone by 4 hours of embryogenesis.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\gd 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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 29 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 6 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of gd
Transgenic constructs containing regulatory region of gd
Deletions and Duplications ( 11 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (0)
No records found.
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (0)
No records found.
Rattus norvegicus (Norway rat) (0)
No records found.
Xenopus tropicalis (Western clawed frog) (0)
No records found.
Danio rerio (Zebrafish) (2)
1 of 15
Yes
Yes
1 of 15
Yes
Yes
Caenorhabditis elegans (Nematode, roundworm) (2)
1 of 15
Yes
No
1 of 15
Yes
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) ( EOG0919066U )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila simulans
Drosophila sechellia
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) ( EOG091504E6 )
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) ( None identified )
No non-Dipteran orthologies identified
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( None identified )
No non-Insect Arthropod orthologies identified
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
No non-Arthropod Metazoa orthologies identified
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (37)
3 of 10
3 of 10
2 of 10
2 of 10
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1 of 10
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1 of 10
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1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
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1 of 10
1 of 10
1 of 10
1 of 10
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1 of 10
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1 of 10
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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 ( 0 )
    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.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    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
    esyN Network Diagram
    Show neighbor-neighbor interactions:
    Select Layout:
    Legend:
    Protein
    RNA
    Selected Interactor(s)
    Interactions Browser

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    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)
    Extracellular Spatzle Activating Pathway Core Components -
    In Drosophila, activation of the Toll pathway is controlled by the generation of a cleaved, active, Toll-binding form of spatzle ligand. Proteolytic activation of spatzle ligand lies downstream of several zymogen activation cascades that are initiated by different cues. In dorsal-ventral patterning, localized activation of spz results in ventral nuclear accumulation of dl. During viral and fungal and gram positive bacterial immune challenge, zymogen cascades are activated by extracellular pattern recognition receptors or virulence factor-mediated cleavage of the zymogen persephone (psh). (Adapted from FBrf0091014 and FBrf0223077).
    External Data
    Linkouts
    Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    X
    Recombination map
    1-38
    Cytogenetic map
    Sequence location
    X:11,985,154..11,987,764 [-]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    11A1-11A1
    Limits computationally determined from genome sequence between P{EP}ATP7EP308 and P{EP}EP1451&P{EP}EP1629
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    11A2-11A4
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Notes
    97 of 129 recombinants between m and fw ocurred distal to cac. 5 of 8 recombinants between cac and fw were distal to gd. 2 of 10 gd/fw recombinants were distal to tsg.
    Stocks and Reagents
    Stocks (8)
    Genomic Clones (13)
     

    Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

    cDNA Clones (9)
     

    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: gd CG1505
      Source for database merge of
      Additional comments
      Other Comments
      Interallelic complementation data suggests that there are two discrete functional domains of gd.
      gd appears to catalyse the ventral activation of the proteolytic cascade producing the Tl ligand.
      gd is a member of the serine protease superfamily of genes which also includes snk, ea and ndl, providing evidence for the involvement of a protease cascade in generating an asymmetric signal during establishment of dorsal-ventral polarity in the embryo.
      Some of the proteins of apico-lateral junctions are required both for apico-basal cell polarity and for the signalling mechanisms controlling cell proliferation, whereas others are required more specifically in cell-cell signalling.
      gd, along with snk and ea, appears to be a component of a serine protease cascade that transduces a spatially and temporally regulated ventralising signal in the perivitelline space of the developing embryo.
      Double mutant combinations of gd with ea alleles demonstrate that spatial regulation of ea activity by localized zymogen activation is a key initial event in defining the polarity of the dorsal-ventral embryonic pattern.
      Mutuants cause maternal-effect lethality. Embryos produced by homozygous females exhibit hyperplasia of dorsal cuticular elements and aplasia of ventral elements. Polarity of the egg shell unaffected. Temperature-sensitive period of temperature-sensitive alleles begins several hours prior to oviposition and persists until 1.5 h after fertilization. Five of eight mutants classed as weak on the basis of filzkorper development; these five tend to be more dorsalized in posterior than in anterior parts of the embryo, as the setal bands become progressively narrowed posteriorly.
      In the absence of gd activity in the perivitelline fluid, no Tl ligand is detected suggesting that gd activity is involved in the production of the Tl ligand.
      Strong dorsalizing mutants of ndl, snk, gd or pip have wild type axial ratios in pupae.
      Flies heterozygous for RpII215Ubl and a gd mutation show no reduction in fertility. gd therefore shows no dominant effects.
      Mutations at gd causes dorsalised embryonic phenotypes.
      Gastrula exhibits excessive furrowing both dorsally and ventrally.
      Origin and Etymology
      Discoverer
      Etymology
      Identification
      External Crossreferences and Linkouts ( 41 )
      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
      Other crossreferences
      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.
      Flygut - An atlas of the Drosophila adult midgut
      Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
      GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
      InterPro - A database of protein families, domains and functional sites
      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 (15)
      Reported As
      Secondary FlyBase IDs
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        References (181)