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General Information
Symbol
Dmel\fog
Species
D. melanogaster
Name
folded gastrulation
Annotation Symbol
CG9559
Feature Type
FlyBase ID
FBgn0000719
Gene Model Status
Stock Availability
Gene Snapshot
folded gastrulation (fog) encodes a secreted ligand that signals via GPCRs to co-ordinate cell shape change during gastrulation; in the nervous system, it regulates motor axon guidance, glial organization and morphogenesis. [Date last reviewed: 2019-03-07]
Also Known As
114, l(1)114, l114, M67
Key Links
Genomic Location
Cytogenetic map
Sequence location
X:22,824,317..22,855,194 [-]
Recombination map
1-66
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Group (FlyBase)
Protein Family (UniProt)
-
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
Predictions / Assertions
-
Summaries
Gene Group (FlyBase)
UNCLASSIFIED RECEPTOR LIGANDS -
Receptor ligands are endogenous polypeptides that bind transmembrane cell surface receptors to elicit a cellular response.
Protein Function (UniProtKB)
Coordinates cell shape changes during formation of the ventral furrow and invagination of the posterior midgut primordium, by inducing apical constriction of cells in spatially and temporally defined manners. Could function as a secreted signal to initiate apical constriction by acting as a ligand for an unidentified G protein-coupled receptor, which in turn activates the G protein alpha subunit encoded by concertina, in neighboring cells. Such an intracellular pathway would ultimately induce contraction of the apical actin-myosin network. In the ventral furrow, fog appears to ensure that all the cells initiate constriction within several minutes of each other. In the posterior midgut invagination, fog appears to direct the ordered progression of constriction initiations out from a central region and also to delimit the peripheral extent of this spreading.
(UniProt, P40795)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
fog: folded gastrulation
Hemizygous lethal; the cellular blastoderm apparently normal and gastrulation begins at normal time. No posterior midgut formed; germ band does not elongate onto dorsal side of embryo but it is thrown into a series of transverse ventral folds. Older embryos often twist around the longitudinal axis for one complete turn; many exhibit an anterodorsal hole through which the brain protrudes and a split in the posterior CNS and protrusion of the midgut. Hemizygous deficiency gives same phenotype. No effect of maternal genotype; homozygous germ line clones make eggs capable of supporting normal embryonic development. Fate mapping localizes fog lethal focus to the posterior pole most likely in presumptive posterior-midgut or proctodaeum cells.
Summary (Interactive Fly)
secreted - novel protein - regulates gastrulation via GPCRs - necessary for coordination of cell shape changes throughout the invaginating primordia - regulates motor axon guidance, glial organization and morphogenesis
Gene Model and Products
Number of Transcripts
6
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\fog or the JBrowse view of Dmel\fog 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 5.53
Annotated transcripts do not represent all supported alternative splices within 5' UTR.
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.46
Gene model reviewed during 5.56
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0077342
4647
730
FBtr0100538
3937
730
FBtr0333179
4504
730
FBtr0333180
3188
730
FBtr0333181
3825
730
FBtr0333182
5256
730
Additional Transcript Data and Comments
Reported size (kB)
3.7 (longest cDNA)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0077034
78.2
730
4.10
FBpp0099983
78.2
730
4.10
FBpp0305381
78.2
730
4.10
FBpp0305382
78.2
730
4.10
FBpp0305383
78.2
730
4.10
FBpp0305384
78.2
730
4.10
Polypeptides with Identical Sequences

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

730 aa isoforms: fog-PA, fog-PB, fog-PC, fog-PD, fog-PE, fog-PF
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Post Translational Modification
May be highly O-glycosylated in its Ser/Thr-rich C-terminal part.
(UniProt, P40795)
Crossreferences
InterPro - A database of protein families, domains and functional sites
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\fog 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 (26 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Biological Process (22 terms)
Terms Based on Experimental Evidence (20 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
Cellular Component (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from direct assay
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
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
fog transcripts are detected in the wing disc folds and in the wing pouch.
fog transcript is uniformly expressed in germ band extended embryos. Most CNS glial cells that express fog transcript also express repo.
The pattern of fog transcription precisely precedes the pattern of apical constrictions in the ventral furrow and in the posterior midgut primordia. Zygotic fog transcription begins in the ventral furrow primordium during the beginning of cellularization, about 30 min. before the start of constrictions. The region of fog transcription encompasses a 12-14 cell wide region of the mesoderm primordia. This corresponds to the subset of ventral furrow cells that make up the initial invagination. During invagination, fog transcripts disappear rapidly from the furrow. fog is also expressed in the invagination primordia of the posterior midgut before invagination begins and disappear as the invagination closes.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\fog 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
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 35 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 16 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of fog
Transgenic constructs containing regulatory region of fog
Deletions and Duplications ( 179 )
Disrupted in
Not disrupted in
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) (1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
No
1 of 15
Yes
No
Rattus norvegicus (Norway rat) (2)
1 of 13
Yes
No
1 of 13
Yes
No
Xenopus tropicalis (Western clawed frog) (1)
1 of 12
Yes
No
Danio rerio (Zebrafish) (2)
1 of 15
Yes
No
1 of 15
Yes
No
Caenorhabditis elegans (Nematode, roundworm) (1)
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) ( EOG09190BQH )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
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) ( EOG091502QW )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
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) (0)
No records found.
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
    Interactions Browser
    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
    Pathways
    Gene Group - Pathway Membership (FlyBase)
    External Data
    Linkouts
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    X
    Recombination map
    1-66
    Cytogenetic map
    Sequence location
    X:22,824,317..22,855,194 [-]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    20E2-20F1
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    Experimentally Determined Recombination Data
    Notes
    Stocks and Reagents
    Stocks (14)
    Genomic Clones (23)
    cDNA Clones (33)
     

    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)
      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
       
      polyclonal
      Commercially Available Antibodies
       
      Other Information
      Relationship to Other Genes
      Source for database identify of
      Source for database merge of
      Source for merge of: fog CR41108 CR41109 CR41110
      Additional comments
      Annotations CG9559, CR41108, CR41109 and CR41110 merged as CG9559 (which corresponds to fog) in release 4.3 of the genome annotation.
      Other Comments
      Loss of fog promotes mitochondrial fusion.
      The generation of ectopic midline crossing phenotypes by excess neuron fog signalling requires Ptp52F.
      fog signal is both necessary and sufficient to trigger the relocalisation of myosin to the apical side of the cell.
      New annotation (CR41108) in release 3.2 of the genome annotation. New annotation (CR41109) in release 3.2 of the genome annotation. New annotation (CR41110) in release 3.2 of the genome annotation.
      Activation of the fog/cta pathway results in ectopic cell shape changes in the gastrula. The normal location of the ventral furrow in embryos with uniformly expressed fog suggests the existence of a fog-independent pathway determining mesoderm-specific cell behaviours and invagination. Epistasis experiments indicate this pathway requires sna but not twi expression.
      Uniform fog expression rescues the gastrulation defects of fog null embryos but not cta mutant embryos, arguing that cta acts downstream of fog.
      cad acts in hindgut development through fog, fkh and wg, but does not play a role in activating tll, hkb, byn and bowl which are also required for proper hindgut development.
      The fog gene product is required during gastrulation for two morphogenetic movements, formation of the ventral furrow and invagination of the posterior midgut primordium, and coordinates cell shape changes by inducing apical constriction of cells in a spatially and temporally defined manner. The maternal contribution of fog product can supplement but not substitute for the zygotic activity. Over-expression of fog can induce ectopic constrictions, and hasten the onset of posterior midgut invagination. fog seems to function as a secreted signal that activates the G protein α subunit encoded by cta in neighboring cells.
      In fog mutant embryos the early constricting cells in the ventral furrow are scattered throughout most of the mesoderm primordium. As a result of the reduced number of appropriate cell shape changes the ventral furrow invaginates later than wild type and with a much less regular appearance.
      Muscle phenotype of mutants studied using polarised light microscopy and antibody staining to detect Mhc-lacZ reporter gene expression in muscles.
      Hemizygous lethal; the cellular blastoderm apparently normal and gastrulation begins at normal time. No posterior midgut formed; germ band does not elongate onto dorsal side of embryo but it is thrown into a series of transverse ventral folds. Older embryos often twist around the longitudinal axis for one complete turn; many exhibit an anterodorsal hole through which the brain protrudes and a split in the posterior CNS and protrusion of the midgut. Hemizygous deficiency gives same phenotype. No effect of maternal genotype; homozygous germ line clones make eggs capable of supporting normal embryonic development. Fate mapping localizes fog lethal focus to the posterior pole most likely in presumptive posterior-midgut or proctodaeum cells.
      Zygotically active locus involved in the terminal developmental program in the embryo.
      Mutations in fog block the transition from the first slow stochastic phase of gastrulation to the second rapid phase of constriction in the ventral furrow and the posterior midgut invaginations.
      fog mutants display defective gastrulation, extensive folds in the germ band and ventral holes in the cuticle particularly at the anterior and posterior end.
      Origin and Etymology
      Discoverer
      Etymology
      Identification
      External Crossreferences and Linkouts ( 42 )
      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
      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
      InterPro - A database of protein families, domains and functional sites
      KEGG Genes - Molecular building blocks of life in the genomic space.
      modMine - A data warehouse for the modENCODE project
      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
      Synonyms and Secondary IDs (14)
      Reported As
      Symbol Synonym
      l(1)16-3-190
      l(1)20Ba
      Secondary FlyBase IDs
      • FBan0009559
      • FBgn0069907
      • FBgn0069908
      • FBgn0069909
      • FBtr0077342
      • FBpp0077034
      Datasets (0)
      Study focus (0)
      Experimental Role
      Project
      Project Type
      Title
      References (180)