FB2026_02 , released June 18, 2026
Gene: Dmel\grnd
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General Information
Symbol
Dmel\grnd
Species
D. melanogaster
Name
grindelwald
Annotation Symbol
CG10176
Feature Type
protein_coding_gene
FlyBase ID
FBgn0032682
Gene Model Status
Current
Stock Availability
7 publicly available
Gene Summary
grindelwald (grnd) encodes a receptor of the TNF superfamily ligand encoded by egr, which activates the intracellular JNK pathway. It is involved in apoptosis and neoplastic growth. [Date last reviewed: 2019-09-19] (FlyBase Gene Snapshot)
All Summaries
Key Links
Genomic Location
Cytogenetic map
36F5-36F5
Sequence location
Recombination map
2-53
RefSeq locus
NT_033779 REGION:18485417..18488021
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (9 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables tumor necrosis factor receptor activity
inferred from direct assay
(Palmerini et al., 2021, Andersen et al., 2015)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (5 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
acts_upstream_of negative regulation of reactive oxygen species biosynthetic process
inferred from mutant phenotype
(Muliyil et al., 2020)
involved_in positive regulation of JNK cascade
inferred from mutant phenotype
(Andersen et al., 2015)
involved_in positive regulation of neuron remodeling
inferred from genetic interaction with FLYBASE:bsk; FB:FBgn0000229
(Zheng et al., 2025)
acts_upstream_of positive regulation of programmed cell death
inferred from mutant phenotype
(Palmerini et al., 2021)
inferred from genetic interaction with FLYBASE:egr; FB:FBgn0033483
(Andersen et al., 2015)
involved_in tumor necrosis factor-mediated signaling pathway
inferred from genetic interaction with FLYBASE:bsk; FB:FBgn0000229
(Zheng et al., 2025)
inferred from genetic interaction with FLYBASE:egr; FB:FBgn0033483
(Andersen et al., 2015)
inferred from mutant phenotype
(Palmerini et al., 2021, de Vreede et al., 2018, Agrawal et al., 2016)
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (3 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
is_active_in plasma membrane
inferred from direct assay
(Andersen et al., 2015)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
located_in apical plasma membrane
inferred from experiment
(Palmerini et al., 2021, Andersen et al., 2015)
located_in cytoplasmic vesicle membrane
inferred from experiment
(Palmerini et al., 2021)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
    -
    Summaries
    Gene Snapshot
    grindelwald (grnd) encodes a receptor of the TNF superfamily ligand encoded by egr, which activates the intracellular JNK pathway. It is involved in apoptosis and neoplastic growth. [Date last reviewed: 2019-09-19]
    Pathway (FlyBase)
    TNFalpha-EIGER SIGNALING PATHWAY CORE COMPONENTS -
    The Tumor Necrosis Factor α (TNFα) signaling pathway is activated by Eiger (egr) binding to a member of the TNF receptor superfamily. The signal is primarily transduced via the Jun N-terminal kinase (JNK) cascade, leading to cell death. (Adapted from FBrf0225608.)
    Gene Group (FlyBase)
    TUMOR NECROSIS FACTOR RECEPTORS -
    The tumor necrosis factor receptors (TNFRs) are type I transmembrane proteins that bind tumor necrosis factors (TNFs) via an extracellular cysteine-rich domain. (Adapted from FBrf0162270.)
    Protein Function (UniProtKB)
    Acts as a receptor for TNF-cytokine egr (PubMed:25874673, PubMed:27076079, PubMed:33824334). Plays a role in activation of JNK signaling and is required for egr-induced apoptosis, including in wing imaginal discs during development (PubMed:25874673, PubMed:33824334). May also play an egr-independent role in cell proliferation (PubMed:25874673). TNF receptor involved in triggering JNK-dependent proliferation of the enteroblast-enterocyte lineage in response to stress-induced release of egr by intestinal stem cells and enteroblasts (PubMed:38944657). Involved in regulation of insulin production in response to dietary protein shortage keeping systemic growth in check (PubMed:27076079). Activation in brain insulin producing cells through binding of egr released into the hemolymph in response to dietary amino acid shortage, results in JNK-dependent inhibition of insulin production (PubMed:27076079).
    (UniProt, Q9VJ83)
    Summary (Interactive Fly)

    tumour necrosis factor receptor - JNK signalling - integrates apoptotic functions from Eiger with signaling from apical polarity determinants to induce JNK-dependent cell death or tumour growth

    Gene Model and Products
    Number of Transcripts
    2
    Number of Unique Polypeptides
    1

    Please see the JBrowse view of Dmel\grnd 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
    Structure
    Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q9VJ83)

    If you don't see a structure in the viewer, refresh your browser.
    Model Confidence:
    • Very high (pLDDT > 90)
    • Confident (90 > pLDDT > 70)
    • Low (70 > pLDDT > 50)
    • Very low (pLDDT < 50)

    AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

    Experimentally Determined Structures
    Crossreferences
    PDB - An information portal to biological macromolecular structures
    Comments on Gene Model

    Gene model reviewed during 5.45

    Transcript Data
    Annotated Transcripts
    Name
    FlyBase ID
    RefSeq ID
    Length (nt)
    Assoc. CDS (aa)
    grnd-RB
    FBtr0300674
    NM_136036
    1789
    241
    grnd-RC
    FBtr0332321
    NM_001273609
    2100
    241
    Additional Transcript Data and Comments
    Reported size (kB)
    Comments
    External Data
    Crossreferences
    Polypeptide Data
    Annotated Polypeptides
    Name
    FlyBase ID
    Predicted MW (kDa)
    Length (aa)
    Theoretical pI
    UniProt
    RefSeq ID
    GenBank
    grnd-PB
    FBpp0289898
    26.3
    241
    7.36
    Q9VJ83
    NP_609880
    AAF53671
    grnd-PC
    FBpp0304599
    26.3
    241
    7.36
    Q9VJ83
    NP_001260538
    AGB93073
    Polypeptides with Identical Sequences

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

    241 aa isoforms: grnd-PB, grnd-PC
    Additional Polypeptide Data and Comments
    Reported size (kDa)
    Comments
    External Data
    Subunit Structure (UniProtKB)

    Interacts (via extracellular cysteine-rich domain) with egr (via secreted TNF-homology soluble form); forms heterohexamers when 3 copies associate with egr trimers (PubMed:25874673, PubMed:33824334). Interacts with Traf6/TRAF2 and veli (via PDZ domain) (PubMed:25874673).

    (UniProt, Q9VJ83)
    Post Translational Modification

    N-glycosylated on Asn-63 (PubMed:29870719). Glycosylation regulates ligand binding, specifically reducing affinity for the TNF egr, thereby inhibiting activation of JNK signaling (PubMed:29870719, PubMed:38944657).

    (UniProt, Q9VJ83)
    Crossreferences
    PDB - An information portal to biological macromolecular structures
    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\grnd using the Feature Mapper tool.

    External Data
    Crossreferences
    Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
    Linkouts
    Expression Data
    Testis-specificity index

    The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

    -0.68

    Transcript Expression
    Polypeptide Expression
    Additional Descriptive Data
    Marker for
     
    Subcellular Localization
    CV Term
    Evidence
    References
    is_active_in plasma membrane
    inferred from direct assay
    (Andersen et al., 2015)
    Expression Deduced from Reporters
    High-Throughput Expression Data
    Associated Tools

    JBrowse - Visual display of RNA-Seq signals

    View Dmel\grnd in JBrowse
    RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
    Bulk Downloads
    RNA-Seq RPKM values for all genes
    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
    DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
    EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
    FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
    FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
    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
    FlyBase Wiki
    Image Based Resources
    Alleles, Insertions, Transgenic Constructs, and Aberrations
    Classical and Insertion Alleles ( 10 )
    For All Classical and Insertion Alleles Show
     
    Other relevant insertions
    Transgenic Constructs ( 12 )
    For All Alleles Carried on Transgenic Constructs Show
    Transgenic constructs containing/affecting coding region of grnd
    Transgenic constructs containing regulatory region of grnd
    Aberrations (Deficiencies and Duplications) ( 6 )
    Variants
    Variant Molecular Consequences
    Alleles Representing Disease-Implicated Variants
    Phenotypes
    Orthologs
    Downloads
    Download All DIOPT Orthologs
    Human Orthologs (via DIOPT v9.1)
    Species\Gene Symbol
    Score
    Best Score
    Best Reverse Score
    Source
    Compara
    Domainoid
    eggNOG
    Hieranoid
    Homologene
    Inparanoid
    OMA
    OrthoDB
    OrthoFinder
    OrthoInspector
    orthoMCL
    Panther
    Phylome
    SonicParanoid
    Alignment
    Complementation?
    Transgene?
    Homo sapiens (Human) (0)
    Model Organism Orthologs (via DIOPT v9.1)
    Species\Gene Symbol
    Score
    Best Score
    Best Reverse Score
    Source
    Compara
    Domainoid
    eggNOG
    Hieranoid
    Homologene
    Inparanoid
    OMA
    OrthoDB
    OrthoFinder
    OrthoInspector
    orthoMCL
    Panther
    Phylome
    SonicParanoid
    Alignment
    Complementation?
    Transgene?
    Rattus norvegicus (Norway rat) (0)
    Mus musculus (laboratory mouse) (0)
    Xenopus tropicalis (Western clawed frog) (0)
    Danio rerio (Zebrafish) (0)
    Caenorhabditis elegans (Nematode, roundworm) (0)
    Anopheles gambiae (African malaria mosquito) (1)
    Agam\AgaP_AGAP008399
    5 of 12
    Yes
    Yes
    Arabidopsis thaliana (thale-cress) (0)
    Saccharomyces cerevisiae (Brewer's yeast) (0)
    Schizosaccharomyces pombe (Fission yeast) (0)
    Escherichia coli (enterobacterium) (0)
    Other Organism Orthologs (via OrthoDB)
    Data provided directly from OrthoDB:grnd. Refer to their site for version information.
    Paralogs
    Downloads
    Download All DIOPT Paralogs
    Paralogs (via DIOPT v9.1)
    Human Disease Associations
    FlyBase Human Disease Model Reports
    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 ( 5 )
    Disease Associations of Human Orthologs (via DIOPT v9.1 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
    Summary of Genetic Interactions
    Interaction Browsers
    View in FlyBase Interactions Browser   View in MIST tool (external link)
    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    grnd
    suppressible
    bsk
    (Zheng et al., 2025)
    grnd
    suppressible
    hep
    (Andersen et al., 2015)
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Alg3
    suppressible
    grnd
    (de Vreede et al., 2018)
    Alg9
    suppressible
    grnd
    (de Vreede et al., 2018)
    crb
    suppressible
    grnd
    (Andersen et al., 2015)
    egr
    suppressible
    grnd
    (Zheng et al., 2025, Andersen et al., 2015)
    HUWE1
    suppressible
    grnd
    (Yanku et al., 2018)
    Noc1
    suppressible
    grnd
    (Mirzoyan et al., 2023)
    Pi3K92E|btl::Egfr
    suppressible
    grnd
    (Portela et al., 2019)
    Ras85D|dlg1
    suppressible
    grnd
    (Andersen et al., 2015)
    Ras85D|scrib
    suppressible
    grnd
    (Andersen et al., 2015)
    RpS3|yki
    suppressible
    grnd
    (Wada et al., 2021)
    scrib
    suppressible
    grnd
    (Andersen et al., 2015)
    Syx7
    suppressible
    grnd
    (Andersen et al., 2015)
    wgn
    enhanceable
    grnd
    (Zheng et al., 2025)
    External Data
    Subunit Structure (UniProtKB)
    Interacts (via extracellular cysteine-rich domain) with egr (via secreted TNF-homology soluble form); forms heterohexamers when 3 copies associate with egr trimers (PubMed:25874673, PubMed:33824334). Interacts with Traf6/TRAF2 and veli (via PDZ domain) (PubMed:25874673).
    (UniProt, Q9VJ83 )
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    FlyBase
    External Links
    External Data
    Linkouts
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2L
    Recombination map
    2-53
    Cytogenetic map
    36F5-36F5
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    36F5-36F5
    Limits computationally determined from genome sequence between P{lacW}Aac11k06710 and P{EP}CG10413EP2164
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    Experimentally Determined Recombination Data
    Location

    2-53

    (FlyBase, 2016)

    2-53.3

    (Comeron et al., 2012)
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (7)
    Genomic Clones (19)
     

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

    cDNA Clones (25)
     

    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 JBrowse for alignment of the cDNAs and ESTs to the gene model.

    cDNA clones, fully sequenced
    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
      Other clones
      RNAi and Array Information
      Linkouts
      DRSC - Results frm RNAi screens
      Antibody Information
      Laboratory Generated Antibodies
       

      monoclonal

      (de Vreede et al., 2018)

      polyclonal

      (Andersen et al., 2015)
      Commercially Available Antibodies
       
      Cell Line Information
      Publicly Available Cell Lines
       
        Other Stable Cell Lines
         
          Other Comments

          dsRNA has been made from templates generated with primers directed against this gene.

          (Pilot et al., 2006)
          Relationship to Other Genes
          Source for database merge of
          Additional comments
          Nomenclature History
          Source for database identify of

          Source for identity of: grnd CG10176

          (Andersen et al., 2015)
          Nomenclature comments
          Etymology

          Named 'grindelwald' ('grnd'), after a village at the foot of Eiger, a Swiss mountain that lent its name to the unique Drosophila TNF, 'Egr'.

          (Andersen et al., 2015)
          Synonyms and Secondary IDs (5)
          Reported As
          Symbol Synonym
          CG10176
          (National Institute of Genetics Fly Stocks, 2022-, Andersen et al., 2015, Nawaz et al., 2012, Biehs et al., 2010, Keleman et al., 2009.8.5, Schnorrer et al., 2007, Pilot et al., 2006, Grad et al., 2004)
          Grnd
          (Hounsell and Fan, 2025, Scopelliti et al., 2025, Kodra et al., 2024, Colombani and Andersen, 2023, Khalili et al., 2023, Havula et al., 2022, Yu et al., 2022, Dillard et al., 2021, Herrera and Bach, 2021, Meschi and Delanoue, 2021, Wada et al., 2021, Fujisawa et al., 2020, La Marca and Richardson, 2020, Portela et al., 2020, Texada et al., 2020, Yanku et al., 2018, Diwanji and Bergmann, 2017)
          grindelwald
          (Agrawal et al., 2016)
          grnd
          (Cruz et al., 2026, Alvarez and Papagiannouli, 2025, Braun et al., 2025, Zheng et al., 2025, Kinoshita et al., 2024, Soares et al., 2024, Wang et al., 2024, Loudhaief et al., 2023, Mirzoyan et al., 2023, Everetts et al., 2021, Klemm et al., 2021, Palmerini et al., 2021, Pathak and Varghese, 2021, Snigdha et al., 2021, Tatapudy et al., 2021, Banreti and Meier, 2020, Krautz et al., 2020, Muliyil et al., 2020, Chang et al., 2019, Hudry et al., 2019, Sanchez et al., 2019, Beira et al., 2018, de Vreede et al., 2018, Gene Disruption Project members, 2018-, Paul et al., 2018, Muzzopappa et al., 2017, Agrawal et al., 2016, Fogarty et al., 2016, Andersen et al., 2015, Bischof and FlyORF project members, 2014.6.20)
          Name Synonyms
          Grindelwald
          (Hounsell and Fan, 2021, Mirzoyan et al., 2019, Paul et al., 2018, Kawamoto et al., 2016)
          grindelwald
          (Fogarty et al., 2016, Andersen et al., 2015)
          Secondary FlyBase IDs
            Datasets (0)
            Study focus (0)
            Experimental Role
            Project
            Project Type
            Title
            Study result (0)
            Result
            Result Type
            Title
            External Crossreferences and Linkouts ( 35 )
            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/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
            UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
            Other crossreferences
            AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
            BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
            DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
            EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
            FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
            FlyMine - An integrated database for Drosophila genomics
            KEGG Genes - Molecular building blocks of life in the genomic space.
            MARRVEL_MODEL - MARRVEL (model organism gene)
            PDB - An information portal to biological macromolecular structures
            Linkouts
            BioGRID - A database of protein and genetic interactions.
            Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
            DroID - A comprehensive database of gene and protein interactions.
            DRSC - Results frm RNAi screens
            Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
            FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
            FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
            Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
            Flygut - An atlas of the Drosophila adult midgut
            iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
            Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
            MIST (protein-protein) - An integrated Molecular Interaction Database
            References (97)