FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\PNUTS
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General Information
Symbol
Dmel\PNUTS
Species
D. melanogaster
Name
Phosphatase 1 nuclear targeting subunit
Annotation Symbol
CG33526
Feature Type
protein_coding_gene
FlyBase ID
FBgn0053526
Gene Model Status
Current
Stock Availability
10 publicly available
Gene Summary
Phosphatase 1 nuclear targeting subunit (PNUTS) encodes a protein phosphatase 1 binding protein involved in developmental growth and gene expression. [Date last reviewed: 2019-09-26] (FlyBase Gene Snapshot)
All Summaries
Key Links
Genomic Location
Cytogenetic map
21E2-21E2
Sequence location
Recombination map
2-1.5
RefSeq locus
NT_033779 REGION:870465..877038
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 (7 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables protein phosphatase 1 binding
inferred from physical interaction with FLYBASE:flw; FB:FBgn0000711
(Bennett et al., 2006)
inferred from physical interaction with FLYBASE:Pp1-13C; FB:FBgn0003132
(Bennett et al., 2006)
inferred from physical interaction with FLYBASE:Pp1α-96A; FB:FBgn0003134
(Bennett et al., 2006)
inferred from physical interaction with FLYBASE:Pp1-87B; FB:FBgn0004103
(Bennett et al., 2006)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables metal ion binding
inferred from electronic annotation with InterPro:IPR000571, InterPro:IPR036855
(InterPro Project Members, 2004-)
enables protein phosphatase 1 binding
inferred from biological aspect of ancestor with PANTHER:PTN005310620
(GO Reference Genome Project, 2011-)
Biological Process (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
involved_in developmental growth
inferred from mutant phenotype
(Ciurciu et al., 2013)
involved_in gene expression
inferred from mutant phenotype
(Ciurciu et al., 2013)
involved_in termination of RNA polymerase II transcription
inferred from mutant phenotype
(Liu et al., 2025)
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (2 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in chromatin
inferred from biological aspect of ancestor with PANTHER:PTN005310620
(GO Reference Genome Project, 2011-)
part_of PTW/PP1 phosphatase complex
inferred from biological aspect of ancestor with PANTHER:PTN005310620
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
Phosphatase 1 nuclear targeting subunit (PNUTS) encodes a protein phosphatase 1 binding protein involved in developmental growth and gene expression. [Date last reviewed: 2019-09-26]
Gene Group (FlyBase)
PTW-PP1 COMPLEX FLW VARIANT -
A nuclear protein serine/threonine phosphatase complex that regulates gene expression. The core of the complex is formed by a protein phosphatase 1 catalytic subunit and the scaffolding/nuclear-targeting subunit PNUTS. (Adapted from FBrf0262508 and PMID:20516061.)
PTW-PP1 COMPLEX PP1-13C VARIANT -
A nuclear protein serine/threonine phosphatase complex that regulates gene expression. The core of the complex is formed by a protein phosphatase 1 catalytic subunit and the scaffolding/nuclear-targeting subunit PNUTS. (Adapted from FBrf0262508 and PMID:20516061.)
PTW-PP1 COMPLEX PP1ALPHA-96A VARIANT -
A nuclear protein serine/threonine phosphatase complex that regulates gene expression. The core of the complex is formed by a protein phosphatase 1 catalytic subunit and the scaffolding/nuclear-targeting subunit PNUTS. (Adapted from FBrf0262508 and PMID:20516061.)
PTW-PP1 COMPLEX PP1-87B VARIANT -
A nuclear protein serine/threonine phosphatase complex that regulates gene expression. The core of the complex is formed by a protein phosphatase 1 catalytic subunit and the scaffolding/nuclear-targeting subunit PNUTS. (Adapted from FBrf0262508 and PMID:20516061.)
Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
2

Please see the JBrowse view of Dmel\PNUTS 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 Q7KU01)

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

Annotated transcripts do not represent all possible combinations of alternative exons and/or alternative promoters.

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

Gene model reviewed during 5.45

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
PNUTS-RA
FBtr0091486
NM_001014454
2731
628
PNUTS-RB
FBtr0091487
NM_001014455
2639
628
PNUTS-RD
FBtr0091489
NM_001014453
4818
1135
PNUTS-RE
FBtr0332000
NM_001272921
4030
1135
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
PNUTS-PA
FBpp0077686
66.6
628
10.60
Q9VPU9
NP_001014454
AAF51439
PNUTS-PB
FBpp0077684
66.6
628
10.60
Q9VPU9
NP_001014455
AAF51438
PNUTS-PD
FBpp0077687
120.6
1135
9.86
Q7KU01
NP_001014453
AAN10491
PNUTS-PE
FBpp0304322
120.6
1135
9.86
Q7KU01
NP_001259850
AGB92387
Polypeptides with Identical Sequences

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

628 aa isoforms: PNUTS-PA, PNUTS-PB
1135 aa isoforms: PNUTS-PD, PNUTS-PE
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
UniProt
Mapped Features

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

-1.54

Transcript Expression
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\PNUTS 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
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 9 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 18 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of PNUTS
Transgenic constructs containing regulatory region of PNUTS
Aberrations (Deficiencies and Duplications) ( 1 )
Inferred from experimentation ( 1 )
Gene disrupted in
Df(2L)Exel6003
(Ryder, 2004.4.26)
Inferred from location ( 5 )
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
lethal
lethal - all die before end of larval stage
lethal - all die before end of larval stage (with Df(2L)ast4)
lethal - all die before end of larval stage (with PNUTS9B)
lethal - all die before end of larval stage (with PNUTS13B)
lethal - all die before end of larval stage | recessive
lethal - all die before end of pupal stage, with Scer\GAL4pnr-MD237
lethal - all die during P-stage, with Scer\GAL4Mef2.PR
partially lethal, with Scer\GAL4tin.CΔ4
partially lethal - majority die, with Scer\GAL4pnr-MD237
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
Sterility
Allele
semi-fertile, with PNUTS13B
semi-fertile, with PNUTSEED.EGFP
semi-sterile | somatic clone - tissue specific, with PNUTS13B
semi-sterile | somatic clone - tissue specific, with PNUTSFRT.EGFP
Other Phenotypes
Allele
abnormal cell growth
some die during embryonic stage | parental effect, with PNUTS13B
some die during embryonic stage | parental effect, with PNUTSEED.EGFP
some die during embryonic stage | parental effect | somatic clone - tissue specific, with PNUTS13B
some die during embryonic stage | parental effect | somatic clone - tissue specific, with PNUTSFRT.EGFP
some die during pupal stage, with Scer\GAL4pnr-MD237
Phenotype manifest in
Allele
eye
nucleus | female limited, with PNUTS13B
nucleus | female limited, with PNUTSED.EGFP
nucleus | female limited, with PNUTSEED.EGFP
nucleus | female limited | somatic clone - tissue specific, with PNUTS13B
nucleus | female limited | somatic clone - tissue specific, with PNUTSFRT.EGFP
nurse cell, with PNUTS13B
nurse cell, with PNUTSED.EGFP
nurse cell, with PNUTSEED.EGFP
nurse cell | germline clone | somatic clone - tissue specific, with PNUTS13B
nurse cell | germline clone | somatic clone - tissue specific, with PNUTSFRT.EGFP
ommatidium
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) (2)
Hsap\PPP1R10
10 of 14
Yes
Yes
Hsap\ATXN2
1 of 14
No
No
12  
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) (2)
Rnor\Ppp1r10
9 of 14
Yes
Yes
Rnor\Atxn2
1 of 14
No
No
Mus musculus (laboratory mouse) (2)
Mmus\Ppp1r10
9 of 14
Yes
Yes
Mmus\Atxn2
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (3)
Xtro\ppp1r10
4 of 13
Yes
Yes
Xtro\atxn2
1 of 13
No
No
Xtro\phtf2
1 of 13
No
No
Danio rerio (Zebrafish) (2)
Drer\ppp1r10
10 of 14
Yes
Yes
Drer\atxn2
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (0)
Anopheles gambiae (African malaria mosquito) (1)
Agam\AgaP_AGAP008794
11 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:PNUTS. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (2)
CG31601
1 of 13
ZC3H18
1 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.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    PPP1R10; protein phosphatase 1 regulatory subunit 10
    10 of 14
    603771
        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
        PNUTS
        suppressible
        RpL27A
        (Ciurciu et al., 2013)
        PNUTS
        suppressible
        Tox4
        (Duncalf et al., 2025)
        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)
        2L
        Recombination map
        2-1.5
        Cytogenetic map
        21E2-21E2
        Sequence location
        FlyBase Computed Cytological Location
        Cytogenetic map
        Evidence for location
        21E2-21E2
        Limits computationally determined from genome sequence between P{lacW}l(2)k09610k09610&P{lacW}Sk09538a and P{PZ}l(2)1068510685&P{lacW}Tango14k00619
        Experimentally Determined Cytological Location
        Cytogenetic map
        Notes
        References
        Experimentally Determined Recombination Data
        Location

        2-1.5

        (FlyBase, 2016)

        2-0.7

        (Comeron et al., 2012)
        Left of (cM)
        Right of (cM)
        Notes
        Stocks and Reagents
        Stocks (10)
        Genomic Clones (12)
         

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

        cDNA Clones (201)
         

        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

              PNUTS is a nuclear protein that colocalises with transcriptionally active RpII215 on salivary gland polytene chromosomes.

              (Ciurciu et al., 2013)

              S2 cells treated with dsRNA generated against this gene show reduced phagocytosis of Candida albicans compared to untreated cells.

              (Stroschein-Stevenson et al., 2006)

              Release 2 annotation CG4124 split into two annotations (CG4124 and CG31657) in release 3 of the genome annotation.

              (FlyBase Genome Annotators, 2002-2003)
              Relationship to Other Genes
              Source for database merge of

              Source for merge of: CG4124 CG31657

              (FlyBase, 1996-)
              Additional comments

              Annotations CG4124 and CG31657 merged as CG33526 (which corresponds to PNUTS) in release 4.1 of the genome annotation.

              (FlyBase Genome Annotators, 2005)

              Source for identity of PNUTS CG4124 was sequence comparison ( date:040310 ).

              (FlyBase, 1996-)

              Source for merge of CG4124 CG31657 was a shared cDNA ( date:030728 ).

              (FlyBase, 1996-)
              Nomenclature History
              Source for database identify of

              Source for identity of: PNUTS CG4124

              (FlyBase, 1996-)
              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 ( 62 )
              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/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
              InterPro - A database of protein families, domains and functional sites
              KEGG Genes - Molecular building blocks of life in the genomic space.
              MARRVEL_MODEL - MARRVEL (model organism gene)
              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)
              MIST (protein-protein) - An integrated Molecular Interaction Database
              References (56)