FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\pll
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General Information
Symbol
Dmel\pll
Species
D. melanogaster
Name
pelle
Annotation Symbol
CG5974
Feature Type
FlyBase ID
FBgn0010441
Gene Model Status
Stock Availability
Enzyme Name (EC)
Gene Summary
Pelle (pll) encodes a tyrosine kinase-like family serine-threonine protein kinase that functions in the Toll pathway, a signaling cascade that plays key roles in various developmental and immune processes, notably embryonic dorso-ventral patterning and regulation of antimicrobial peptide production. [Date last reviewed: 2016-06-30] (FlyBase Gene Snapshot)
Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
3-92
RefSeq locus
NT_033777 REGION:27251131..27253182
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (25 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:Tehao; FB:FBgn0026760
inferred from physical interaction with FLYBASE:Pli; FB:FBgn0025574
inferred from physical interaction with UniProtKB:P22812
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
Biological Process (17 terms)
Terms Based on Experimental Evidence (11 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:tub; FB:FBgn0003882
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:Cka; FB:FBgn0044323
inferred from mutant phenotype
Terms Based on Predictions or Assertions (8 terms)
CV Term
Evidence
References
involved_in apoptotic process
non-traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000701353
traceable author statement
traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000784925
inferred from electronic annotation with InterPro:IPR000488
inferred from biological aspect of ancestor with PANTHER:PTN000701353
traceable author statement
Cellular Component (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
inferred from direct assay
located_in plasma membrane
inferred from direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
is_active_in cytoplasm
inferred from biological aspect of ancestor with PANTHER:PTN000784925
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000784925
is_active_in plasma membrane
inferred from biological aspect of ancestor with PANTHER:PTN000701353
Protein Family (UniProt)
Belongs to the protein kinase superfamily. TKL Ser/Thr protein kinase family. Pelle subfamily. (Q05652)
Catalytic Activity (EC/Rhea)
protein serine/threonine kinase activity
Summaries
Gene Snapshot
Pelle (pll) encodes a tyrosine kinase-like family serine-threonine protein kinase that functions in the Toll pathway, a signaling cascade that plays key roles in various developmental and immune processes, notably embryonic dorso-ventral patterning and regulation of antimicrobial peptide production. [Date last reviewed: 2016-06-30]
Pathway (FlyBase)
POSITIVE REGULATORS OF HIPPO SIGNALING PATHWAY -
The Hippo signaling pathway is an intracellular kinase cascade in which hpo kinase in complex with sav, phosphorylates wts kinase which, in turn, phosphorylates yki transcriptional co-activator leading to its cytosolic retention. Positive regulators of the pathway, enhance the cytosolic retention of yki, negatively regulating tissue growth (Adapted from FBrf0224870).
TOLL-NF-kappaB SIGNALING PATHWAY CORE COMPONENTS -
In Drosophila, the canonical Toll signaling pathway is initiated by the binding of a spatzle ligand to Toll (Tl) or a Toll-like receptor leading to the nuclear localization of the NF-κB (dl or Dif) transcription factor. (Adapted from FBrf0091014 and FBrf0223077).
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).
NON-RECEPTOR TKL KINASES -
Non-receptor tyrosine kinase-like (TKL) kinases are a diverse group of intracellular serine-threonine protein kinases with sequence similarity to tyrosine kinases (TK) but lacking TK-specific motifs.
Protein Function (UniProtKB)
Plays an essential role in the Tl receptor signaling pathway that establishes embryonic dorsoventral polarity; the signal directs import of dl into ventral and ventrolateral nuclei, thereby establishing dorsoventral polarity. Tub recruits pll to the plasma membrane and protein-protein interaction activates pll.
(UniProt, Q05652)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
pll: pelle
Maternal-effect lethal; homozygous females sterile. As in the mutant dl, lethal embryos produced by pll females lack ventral and lateral elements (Anderson and Nusslein-Volhard, 1984). Extreme pll embryos consist of a long, hollow tube with dorsal cuticular structures (Muller-Holtkamp et al., 1985). Dorsalization is also observed in the pattern of ftz stripes in pll embryos (Carroll et al., 1987). Injection of wild-type cytoplasm into pll embryos partially restores the normal embryonic pattern of development (Muller-Holtkamp et al., 1985).
Summary (Interactive Fly)

tyrosine kinase - Toll pathway - dorsal group - involved in signal transduction during induction of dorsal/ventral polarity during early embryogenesis - regulation of antimicrobial peptides.

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

Please see the JBrowse view of Dmel\pll 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
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end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q05652)

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 5.47

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0085179
1931
501
FBtr0334732
1852
501
Additional Transcript Data and Comments
Reported size (kB)

1.9 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0084549
56.2
501
8.87
FBpp0306780
56.2
501
8.87
Polypeptides with Identical Sequences

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

501 aa isoforms: pll-PA, pll-PB
Additional Polypeptide Data and Comments
Reported size (kDa)

501 (aa); 56 (kD)

Comments
External Data
Subunit Structure (UniProtKB)

Interacts (via Death domain) with tub (via Death domain). Interacts with Pellino (Pli).

(UniProt, Q05652)
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\pll 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).

-0.44

Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

pll transcripts are expressed most strongly in early embryos and adult females and are observed at a lower level throughout development.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
inferred from direct assay
located_in plasma membrane
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\pll 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
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
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 34 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 14 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of pll
Transgenic constructs containing regulatory region of pll
Aberrations (Deficiencies and Duplications) ( 18 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
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
axon & eye photoreceptor cell (with pll2)
axon & eye photoreceptor cell (with pll7)
axon & eye photoreceptor cell (with pll25)
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (7)
9 of 14
Yes
Yes
5 of 14
No
Yes
5 of 14
No
Yes
4 of 14
No
Yes
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
Yes
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (6)
7 of 14
Yes
Yes
6 of 14
No
Yes
5 of 14
No
Yes
3 of 14
No
Yes
1 of 14
No
No
1 of 14
No
Yes
Mus musculus (laboratory mouse) (6)
8 of 14
Yes
Yes
6 of 14
No
Yes
5 of 14
No
Yes
2 of 14
No
Yes
1 of 14
No
No
1 of 14
No
Yes
Xenopus tropicalis (Western clawed frog) (8)
5 of 13
Yes
Yes
5 of 13
Yes
Yes
3 of 13
No
Yes
3 of 13
No
Yes
2 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
Yes
Danio rerio (Zebrafish) (5)
9 of 14
Yes
Yes
5 of 14
No
Yes
3 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (2)
11 of 14
Yes
Yes
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (2)
11 of 12
Yes
Yes
2 of 12
No
Yes
Arabidopsis thaliana (thale-cress) (529)
4 of 13
Yes
Yes
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Yes
Yes
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Saccharomyces cerevisiae (Brewer's yeast) (2)
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Schizosaccharomyces pombe (Fission yeast) (1)
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Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:pll. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (1)
2 of 13
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.
    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

    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
    Subunit Structure (UniProtKB)
    Interacts (via Death domain) with tub (via Death domain). Interacts with Pellino (Pli).
    (UniProt, Q05652 )
    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
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    3R
    Recombination map
    3-92
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    97E11-97E11
    Limits computationally determined from genome sequence between P{PZ}sda03884 and P{PZ}btzrL203
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    97F-97F
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (19)
    Genomic Clones (11)
     

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

    cDNA Clones (245)
     

    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)
    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

      When dsRNA constructs are made and transiently transfected into S2 cells in RNAi experiments, a decrease in cytokinetic index is seen.

      dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.

      pll is activated by autophosphorylation.

      Shows particularly robust cycling of transcription in adult heads, as assessed by expression analysis using high density oligonucleotide arrays with probe generated during three 12-point time course experiments over the course of 6 days.

      pll functions to downregulate the signal-dependent relocalisation of tub protein.

      pll is required for muscle development in the embryo.

      Targeting of either tub or pll product to the plasma membrane by myristylation is sufficient to activate the signal transduction pathway that leads to translocation of the dl product. Activated Tl induces a localized recruitment of tub and pll proteins to the plasma membrane.

      The protein kinase activity of the pll protein differentially regulates its interactions with the dl and tub proteins.

      tub, pll, cact and dl form a complex essential for signal transduction.

      Yeast two hybrid assay demonstrates both tub and pll interact with dl. Interactions have also been confirmed in an in vitro binding assay.

      The embryonic regulatory pathway, comprising the gene products between spz and cact (Tl, tub and pll) but not the genes acting upstream or downstream (ea and dl), is involved in the induction of the Drs gene in adults.

      Mutations that affect the synthesis of antimicrobial peptides dramatically lower the resistance of flies to infection.

      Expression of pll enhances the transcriptional activity and nuclear localisation of dl. Tl and pll can functionally interact to enhance dl activity synergistically.

      In cultured cells and embryos cact degradation is dependent on a pll mediated signal, mutation of a putative phosphorylation motif blocks signal responsiveness.

      Interaction of the pll kinase with the membrane associated tub protein is required for transduction of the dorsoventral signal in embryos. pll can function in signal transduction in the absence of tub and cannot act upstream of tub in the signaling pathway.

      The interaction of the pll protein with the tub protein is mediated by the putative regulatory domain of pll and not by the protein kinase catalytic domain.

      Signal-dependent dl phosphorylation is modulated by three intracellular proteins, pll, tub and cact.

      tub acts upstream of pll in the dorsoventral signal transduction pathway. The two-hybrid yeast interaction system demonstrates direct interaction of pll (bait) with tub (prey) to activate a reporter gene, Ecol\lacZ. Combinations pll dl and pll cact did not activate Ecol\lacZ expression.

      pll is not essential for viability but lack of pll gene product decreases the chances of survival to adult stages. Different pll alleles interact with a different spectra of tub alleles. Double mutants of tub and pll show higher zygotic lethality than either single mutant.

      pll acts downstream of Tl indicating that the gene product is part of the machinery that transmits the signal from the Tl receptor to promote nuclear localisation of the dl gene product.

      Genetic and molecular characterization of pll shows that it encodes a protein kinase, predicted to have a serine and threonine specificity. The pll kinase represents a new member of the raf/mos family.

      Double mutant combinations of pll 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.

      Recessive dorsalizing mutants of the dorsal group gene pll have significantly reduced axial ratios in pupae.

      Epistatic relationships exist between dorsalizing maternal effect mutations and "dppHin" alleles.

      Relationship to Other Genes
      Source for database merge of
      Additional comments
      Nomenclature History
      Source for database identify of
      Nomenclature comments
      Etymology
      Synonyms and Secondary IDs (10)
      Reported As
      Symbol Synonym
      pll/IRAK1
      Name Synonyms
      Serine/threonine-protein kinase pelle
      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 ( 66 )
        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
        UniProt/TrEMBL - Automatically annotated and unreviewed 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
        KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
        MIST (protein-protein) - An integrated Molecular Interaction Database
        SignaLink - A signaling pathway resource with multi-layered regulatory networks.
        References (377)