FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\hrg
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General Information
Symbol
Dmel\hrg
Species
D. melanogaster
Name
hiiragi
Annotation Symbol
CG9854
Feature Type
protein_coding_gene
FlyBase ID
FBgn0015949
Gene Model Status
Current
Stock Availability
28 publicly available
Enzyme Name (EC)
polynucleotide adenylyltransferase (2.7.7.19)
Gene Summary
hiiragi (hrg) encodes the canonical poly(A) polymerase required for the nuclear poly(A) tail synthesis at the mRNA 3'-end. hrg product is also involved in cytoplasmic polyadenylation of specific mRNAs during early to mid-oogenesis, through its interaction with the CPEB (cytoplasmic polyadenylation element binding) protein encoded by orb. [Date last reviewed: 2018-09-20] (FlyBase Gene Snapshot)
All Summaries
Also Known As

Pap, poly(A) polymerase, l(2)k07619

Key Links
Genomic Location
Cytogenetic map
56E1-56E1
Sequence location
Recombination map
2-90
RefSeq locus
NT_033778 REGION:19663919..19669832
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 (8 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables poly(A) RNA polymerase activity
inferred from direct assay
(Murata et al., 2001)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
enables nucleotidyltransferase activity
inferred from electronic annotation with InterPro:IPR011068
(InterPro Project Members, 2004-)
enables poly(A) RNA polymerase activity
inferred from sequence or structural similarity with SGD:S000001710
(Mount and Salz, 2000)
inferred from biological aspect of ancestor with PANTHER:PTN000074206
(GO Reference Genome Project, 2011-)
inferred from electronic annotation with InterPro:IPR007012
(InterPro Project Members, 2004-)
inferred from electronic annotation with InterPro:IPR007012, InterPro:IPR014492
(InterPro Project Members, 2004-)
enables RNA binding
inferred from electronic annotation with InterPro:IPR007010, InterPro:IPR011068
(InterPro Project Members, 2004-)
Biological Process (3 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
involved_in cytosolic mRNA polyadenylation
inferred from genetic interaction with FLYBASE:orb; FB:FBgn0004882
(Benoit et al., 2008)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
involved_in co-transcriptional mRNA 3'-end processing, cleavage and polyadenylation pathway
inferred from sequence or structural similarity with UniProtKB:P51003
(Gene Ontology Curators, 2002-)
involved_in RNA 3'-end processing
inferred from electronic annotation with InterPro:IPR011068
(InterPro Project Members, 2004-)
Cellular Component (2 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
located_in nucleoplasm
inferred from sequence or structural similarity with UniProtKB:P51003-1
(Gene Ontology Curators, 2002-)
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000074206
(GO Reference Genome Project, 2011-)
located_in nucleus
inferred from electronic annotation with InterPro:IPR014492
(InterPro Project Members, 2004-)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Catalytic Activity (EC/Rhea)
poly(A) RNA polymerase activity
RNA(n) + ATP = RNA(n)-3'-adenine ribonucleotide + diphosphate (2.7.7.19)
RHEA 11332:
Summaries
Gene Snapshot
hiiragi (hrg) encodes the canonical poly(A) polymerase required for the nuclear poly(A) tail synthesis at the mRNA 3'-end. hrg product is also involved in cytoplasmic polyadenylation of specific mRNAs during early to mid-oogenesis, through its interaction with the CPEB (cytoplasmic polyadenylation element binding) protein encoded by orb. [Date last reviewed: 2018-09-20]
Gene Group (FlyBase)
POLY(A) POLYMERASES -
Poly(A) polymerases (PAPs) catalyze the addition of nontemplated adenylyl residues to the 3' ends of RNA substrates. In eukaryotes, canonical PAPs polyadenylate mRNAs prior to export from the nucleus to the cytoplasm, which is required for the stabilization, export and efficient translation of transcripts. In contrast, non-canonical PAPs add poly(A) tails to a different types of RNAs to stimulate their degradation - similar to the function of bacterial poly(A) tails. PAPs therefore play crucial and diverse roles in the regulation of gene expression. (Adapted from PMID:21956911.)
Summary (Interactive Fly)

involved in both nuclear and cytoplasmic polyadenylation of mRNA - controls the formation of the wing margin

Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
2

Please see the JBrowse view of Dmel\hrg 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 A0A0B4KG96)

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

Stop-codon suppression (UGA) postulated; FBrf0216884

Gene model reviewed during 5.44

Gene model reviewed during 5.51

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
hrg-RA
FBtr0086483
NM_080529
3914
659
hrg-RB
FBtr0086484
NM_166372
3607
659
hrg-RC
FBtr0111010
NM_001043097
3493
659
hrg-RD
FBtr0330248
NM_001274168
4043
827
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
hrg-PA
FBpp0085678
74.5
659
7.09
Q9N6D7
NP_536790
AAF57528
hrg-PB
FBpp0085679
74.5
659
7.09
Q9N6D7
NP_725919
AAM68424
hrg-PC
FBpp0110309
74.5
659
7.09
Q9N6D7
NP_001036562
ABI31107
hrg-PD
FBpp0303281
92.3
827
7.81
A0A0B4KG96
NP_001261097
AGB93629
Polypeptides with Identical Sequences

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

659 aa isoforms: hrg-PA, hrg-PB, hrg-PC
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
Mapped Features

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

-1.10

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\hrg 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) 1-3
  • 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 ( 19 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 15 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of hrg
Transgenic constructs containing regulatory region of hrg
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) (3)
Hsap\PAPOLA
13 of 14
Yes
Yes
1  
Hsap\PAPOLB
13 of 14
Yes
Yes
Hsap\PAPOLG
13 of 14
Yes
Yes
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) (3)
Rnor\Papola
13 of 14
Yes
Yes
Rnor\Papolg
13 of 14
Yes
Yes
Rnor\Papolb
1 of 14
No
Yes
Mus musculus (laboratory mouse) (3)
Mmus\Papola
13 of 14
Yes
Yes
Mmus\Papolg
13 of 14
Yes
Yes
Mmus\Papolb
12 of 14
No
Yes
Xenopus tropicalis (Western clawed frog) (4)
Xtro\papolg
8 of 13
Yes
Yes
Xtro\papola
6 of 13
No
Yes
Xtro\LOC100485335
1 of 13
No
No
Xtro\rpl35a
1 of 13
No
No
Danio rerio (Zebrafish) (2)
Drer\papola
13 of 14
Yes
Yes
Drer\papolg
13 of 14
Yes
Yes
Caenorhabditis elegans (Nematode, roundworm) (3)
Cele\pap-1
13 of 14
Yes
Yes
Cele\pap-2
13 of 14
Yes
Yes
Cele\pap-3
13 of 14
Yes
Yes
Anopheles gambiae (African malaria mosquito) (1)
Agam\AgaP_AGAP013207
12 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (5)
Atha\PAPS1
12 of 13
Yes
Yes
Atha\nPAP
10 of 13
No
Yes
Atha\PAPS2
10 of 13
No
Yes
Atha\PAPS3
4 of 13
No
Yes
Atha\AT3G56320
1 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (2)
Scer\PAP1
12 of 13
Yes
Yes
Scer\PAP2
1 of 13
No
No
Schizosaccharomyces pombe (Fission yeast) (1)
Spom\pla1
10 of 12
Yes
Yes
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:hrg. 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?
    PAPOLA; poly(A) polymerase alpha
    13 of 14
    605553
         
        PAPOLB; poly(A) polymerase beta
        13 of 14
        607436
            PAPOLG; poly(A) polymerase gamma
            13 of 14
            616865
                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
                hrg
                enhanceable
                dx
                (Murata et al., 1996)
                hrg
                suppressible
                H
                (Murata et al., 1996)
                hrg
                enhanceable
                N
                (Murata et al., 1996)
                hrg
                enhanceable
                Ser
                (Murata et al., 1996)
                Starting gene(s)
                Interaction type
                Interacting gene(s)
                Reference
                dx
                enhanceable
                hrg
                (Murata et al., 1996)
                H
                suppressible
                hrg
                (Murata et al., 1996)
                N
                enhanceable
                hrg
                (Murata et al., 1996)
                Nab2
                enhanceable
                hrg
                (Pak et al., 2011)
                orb
                enhanceable
                hrg
                (Benoit et al., 2008, Juge et al., 2002)
                Ser
                enhanceable
                hrg
                (Murata et al., 1996)
                External Data
                Linkouts
                BioGRID - A database of protein and genetic interactions.
                DroID - A comprehensive database of gene and protein interactions.
                MIST (genetic) - An integrated Molecular Interaction Database
                MIST (protein-protein) - An integrated Molecular Interaction Database
                Pathways
                Signaling Pathways (FlyBase)
                Metabolic Pathways
                FlyBase
                External Links
                External Data
                Linkouts
                KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
                Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                Class of Gene
                Genomic Location and Detailed Mapping Data
                Chromosome (arm)
                2R
                Recombination map
                2-90
                Cytogenetic map
                56E1-56E1
                Sequence location
                FlyBase Computed Cytological Location
                Cytogenetic map
                Evidence for location
                56E1-56E1
                Limits computationally determined from genome sequence between P{PZ}sm05338&P{PZ}emm1 and P{lacW}l(2)k08002k08002
                Experimentally Determined Cytological Location
                Cytogenetic map
                Notes
                References
                56E-56E
                (Murata, 1998.10.6)
                56D11-56E6
                (Juge, 2000.2.4)
                43E15-43E16
                (determined by in situ hybridisation)
                (Spradling et al., 1999)
                56E-56E
                (determined by in situ hybridisation)
                (Murata et al., 1996)
                43E15-43E16
                (BDGP Project Members, 1994-1999)
                Experimentally Determined Recombination Data
                Location

                2-90

                (FlyBase, 2016)

                2-95.0

                (Comeron et al., 2012)
                Left of (cM)
                (Murata et al., 1996)
                Right of (cM)
                (Murata et al., 1996)
                Notes
                Stocks and Reagents
                Stocks (28)
                Genomic Clones (18)
                 

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

                cDNA Clones (62)
                 

                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

                (Na et al., 2016)
                Commercially Available Antibodies
                 
                Cell Line Information
                Publicly Available Cell Lines
                 
                  Other Stable Cell Lines
                   
                    Other Comments

                    Regulation of hrg level is essential for controlled cytoplasmic polyadenylation and early development.

                    (Juge et al., 2002)

                    Identification: By sequence similarity at low stringency to a bovine PAP cDNA.

                    (Simonelig et al., 2000)

                    Strong mutants are lethal at the late embryonic stage.

                    (Simonelig et al., 2000)

                    hrg is involved in the Ser system, rather that the Dl system.

                    (Murata et al., 1996)

                    Mutant alleles show wing scalloping.

                    (Murata et al., 1996)

                    hrg is involved in the N signalling cascade induced by Ser rather than Dl.

                    (Murata et al., 1996)
                    Relationship to Other Genes
                    Source for database merge of

                    Source for merge of: hrg Pap

                    (Murata et al., 2001)

                    Source for merge of: hrg l(2)k07619

                    (Mutsuddi et al., 2004)

                    Source for merge of: Pap CG9854

                    (FlyBase, 1996-)
                    Additional comments

                    Source for merge of Pap CG9854 was sequence comparison ( date:000725 ).

                    (FlyBase, 1996-)
                    Nomenclature History
                    Source for database identify of
                    Nomenclature comments
                    Etymology
                    Synonyms and Secondary IDs (12)
                    Datasets (0)
                    Study focus (0)
                    Experimental Role
                    Project
                    Project Type
                    Title
                    Study result (0)
                    Result
                    Result Type
                    Title
                    External Crossreferences and Linkouts ( 72 )
                    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)
                    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 (genetic) - An integrated Molecular Interaction Database
                    MIST (protein-protein) - An integrated Molecular Interaction Database
                    Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                    References (91)