FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\ndl
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General Information
Symbol
Dmel\ndl
Species
D. melanogaster
Name
nudel
Annotation Symbol
CG10129
Feature Type
FlyBase ID
FBgn0002926
Gene Model Status
Stock Availability
Enzyme Name (EC)
Gene Summary
nudel (ndl) encodes a multi-functional protein with serine protease activity. It is required for both eggshell biogenesis and embryonic dorsoventral patterning. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
Also Known As

SP89

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
3-17
RefSeq locus
NT_037436 REGION:6593938..6603366
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (11 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
Biological Process (8 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
involved_in egg activation
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
involved_in protein processing
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
involved_in proteolysis
inferred from electronic annotation with InterPro:IPR001254, InterPro:IPR018114
inferred from sequence model
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)
Protein Family (UniProt)
Belongs to the peptidase S1 family. (P98159)
Catalytic Activity (EC/Rhea)
serine-type endopeptidase activity
Summaries
Gene Snapshot
nudel (ndl) encodes a multi-functional protein with serine protease activity. It is required for both eggshell biogenesis and embryonic dorsoventral patterning. [Date last reviewed: 2019-03-14]
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 - TRYPSIN-LIKE -
Trypsin-like serine proteases of the S1A family are endopeptidases that cleave amide substrates following Arginine or Lysine 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:7671306). A protease cascade involving ndl, gd, snk and ea results in activation of the spz Toll receptor ligand; acts upstream of gd, snk and ea and is required for proteolytic processing of gd (PubMed:20605458). Activation of ea requires activation of the ndl-gd-snk protease cascade and sulfation of a vitelline membrane component by pip (PubMed:20605458). Localized activation of the Toll receptor in the ventral region of the embryo defines cell identities along the dorsal-ventral continuum (Probable).
(UniProt, P98159)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
ndl: nudel
Maternal effect mutation producing totally dorsalized embryos. Spacing of transverse stripes of ftz protein altered in mutant embryos (Carroll, Winslow, Twombly, and Scott, 1987, Development 99: 327-32).
Summary (Interactive Fly)

serine protease - docking protein - dorsal group - part of a signaling cascade that determines dorsal/ventral polarity of the embryo - required for eggshell biogenesis

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

Please see the JBrowse view of Dmel\ndl 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
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end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry P98159)

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
Comments on Gene Model

Gene model reviewed during 5.46

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0076984
8230
2616
Additional Transcript Data and Comments
Reported size (kB)

8.4 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0076693
292.5
2616
5.77
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)

2616 (aa); 292 (kD predicted)

Comments
External Data
Post Translational Modification

Requires cleavage for activation (presumably).

(UniProt, P98159)
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\ndl 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).

-0.34

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

ndl transcripts are expressed almost exclusively in follicle cells. They are first detected at stage S7 of oogenesis but reach a peak at stages S9 and S10. In the majority of egg chambers, the transcript is asymmetrically distributed with a higher level seen in ventral than in dorsal follicle cells. The transcript decreases in abundance dramatically in late stage S10 and is undetectable by stage S11.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data

Proteolytically processed segments of ndl protein are secreted into the perivitelline space and stably localized.

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

JBrowse - Visual display of RNA-Seq signals

View Dmel\ndl in JBrowse
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
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
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 23 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 10 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of ndl
Transgenic constructs containing regulatory region of ndl
Aberrations (Deficiencies and Duplications) ( 3 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (29)
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Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (29)
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Mus musculus (laboratory mouse) (30)
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Xenopus tropicalis (Western clawed frog) (37)
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Danio rerio (Zebrafish) (39)
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Caenorhabditis elegans (Nematode, roundworm) (7)
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Anopheles gambiae (African malaria mosquito) (17)
10 of 12
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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:ndl. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (39)
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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 ( 0 )
    Allele
    Disease
    Interaction
    References
    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
    Interaction Browsers

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    Interaction Browsers
    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.
    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)
    3L
    Recombination map
    3-17
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    65B5-65B5
    Limits computationally determined from genome sequence between P{PZ}l(3)0209402094 and P{lacW}l(3)L4060L4060
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    Experimentally Determined Recombination Data
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (8)
    Genomic Clones (23)
    cDNA Clones (6)
     

    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
               
              Commercially Available Antibodies
               
              Cell Line Information
              Publicly Available Cell Lines
               
                Other Stable Cell Lines
                 
                  Other Comments

                  The non-protease domains of ndl do not influence the signaling pathway that establishes embryonic DV polarity.

                  ndl has an integral role in the biogenesis of the vitelline membrane and may function in embryonic dorsal ventral patterning through creation of a matrix structure necessary for activity of the dorsal ventral protease cascade.

                  Catalytic activity of the protease domain of the modular ndl protein is required for dorsoventral patterning of the embryo.

                  The serine protease activity of ndl protein is essential for embryonic dorsoventral polarity. Active ndl protease is generated by autoproteolytic cleavage of a zymogen form. This activation is independent of the other known proteases involved in establishment of dorsoventral polarity and appears to occur symmetrically on the surface of the embryo.

                  A follicular cell marker system that yields a visible phenotype within the mature egg shell allows direct comparison of a clone and its effect on the dorsal ventral pattern of the embryos. Follicular epithelium has a lack of a localised requirement for ndl protease activity.

                  Phenotypic and genetic analysis reveals that ndl has two distinct roles important for embryogenesis: one in establishing dorsoventral polarity, the other in ensuring structural integrity of the egg.

                  ndl is a somatic-dependent gene required to activate and localise the serine protease cascade that produces Tls ligand.

                  Local activation of the protease cascade within the extra embryonic perivitelline compartment that defines embryonic dorsoventral polarity depends on a positional cue that is laid down during oogenesis outside the oocyte, ndl encodes an essential component of this cue. ndl function is required during oogenesis and also during the early stages of embryogenesis.

                  Double mutant combinations of ndl 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.

                  Strong dorsalizing mutants of ndl, snk, gd or pip have wild type axial ratios in pupae.

                  Analysis of mosaic females indicates that the expression of ndl, pip and wbl is required in the somatic tissue, in the follicle cells that surround the oocyte, for the production of embryos with the correct dorsoventral axis.

                  Relationship to Other Genes
                  Source for database merge of
                  Additional comments
                  Nomenclature History
                  Source for database identify of

                  Source for identity of: ndl CG10129

                  Nomenclature comments
                  Etymology
                  Synonyms and Secondary IDs (9)
                  Datasets (0)
                  Study focus (0)
                  Experimental Role
                  Project
                  Project Type
                  Title
                  Study result (0)
                  Result
                  Result Type
                  Title
                  External Crossreferences and Linkouts ( 34 )
                  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.
                  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)
                  MEROPS - An information resource for peptidases (also termed proteases, proteinases and proteolytic enzymes) and the proteins that inhibit them.
                  Linkouts
                  BioGRID - A database of protein and genetic interactions.
                  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
                  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 (154)