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⇱ FlyBase Gene Report: Dmel\bol

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FB2026_01 , released March 12, 2026
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General Information
Symbol
Dmel\bol
Species
D. melanogaster
Name
boule
Annotation Symbol
CG4760
Feature Type
FlyBase ID
FBgn0011206
Gene Model Status
Stock Availability
Gene Summary
boule (bol) encodes a translational regulator required in spermatogenesis for entry into meiosis and spermatid differentiation. It has an additional role in mushroom body γ neurons as a negative regulator of axon pruning. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
Also Known As

lincRNA.485

Genomic Location
Cytogenetic map
Sequence location
Recombination map
3-28
RefSeq locus
NT_037436 REGION:9097629..9134405
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (14 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables RNA binding
inferred from mutant phenotype
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN008962316
enables mRNA binding
inferred from sequence or structural similarity
inferred from electronic annotation with InterPro:IPR035979
enables RNA binding
inferred from electronic annotation with InterPro:IPR000504
inferred from biological aspect of ancestor with PANTHER:PTN008513653
Biological Process (6 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:twe; FB:FBgn0002673
inferred from mutant phenotype
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN008513653
Cellular Component (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
located_in nucleus
inferred from direct assay
located_in perikaryon
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in cytoplasm
inferred from biological aspect of ancestor with PANTHER:PTN008513653
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the RRM DAZ family. (Q24207)
Protein Signatures (InterPro)
Summaries
Gene Snapshot
boule (bol) encodes a translational regulator required in spermatogenesis for entry into meiosis and spermatid differentiation. It has an additional role in mushroom body γ neurons as a negative regulator of axon pruning. [Date last reviewed: 2019-03-07]
(Alliance, FBgn0011206 )
Protein Function (UniProtKB)
RNA-binding protein that plays a central role in spermatogenesis. Required for meiotic entry and germline differentiation, at the transition between G2 and M phases of meiosis I. Acts by regulating translation of specific mRNAs, possibly by binding to their 3'-UTR. Essential for translation of twine (twe) mRNA. Required for the expression of various genes such as CG6784, CG17210, CG15841 scpr-B, scpr-C, and rho-6.
(UniProt, Q24207)
Summary (Interactive Fly)

RRM motif protein involved in spermatogenesis - a translational regulator - required in spermatogenesis for entry into meiosis and spermatid differentiation - a negative regulator of axon pruning in mushroom body neurons

Gene Model and Products
Number of Transcripts
9
Number of Unique Polypeptides
6

Please see the JBrowse view of Dmel\bol 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 Q24207)

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

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.55

A subset of the transcripts annotated for this gene are postulated to be circular mRNAs (FBrf0247226).

Gene model reviewed during 6.46

Gene model includes transcripts encoding non-overlapping portions of the full CDS.

Gene model reviewed during 5.51

Gene model reviewed during 5.46

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

Gene model reviewed during 5.45

Stage-specific extension of 3' UTRs observed during embryogenesis (FBrf0215804); all variants may not be annotated.

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

Gene model reviewed during 5.53

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0076541
1801
189
FBtr0076538
2511
228
FBtr0076539
2294
189
FBtr0076540
2360
228
FBtr0300462
2023
95
FBtr0331858
6538
233
FBtr0344979
2930
228
FBtr0481739
626
118
FBtr0481740
303
49
Additional Transcript Data and Comments
Reported size (kB)

3.0 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0076268
20.2
189
8.83
FBpp0076265
24.7
228
7.65
FBpp0076266
20.2
189
8.83
FBpp0076267
24.7
228
7.65
FBpp0304242
25.2
233
7.14
FBpp0311234
24.7
228
7.65
FBpp0428496
12.4
118
8.91
FBpp0428497
5.5
49
3.75
Polypeptides with Identical Sequences

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

228 aa isoforms: bol-PB, bol-PD, bol-PG
189 aa isoforms: bol-PA, bol-PC
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

Interacts with the translational regulator orb2.

(UniProt, Q24207)
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\bol 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).

2.92

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: extended 3' UTR isoform

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

Zygotic-specific isoforms of bol with long 3' UTR extensions were observed.

bol mRNA is lost from the mushroom body gamma neurons by 0 hours APF.

bol transcripts are expressed only in males where they are germline specific.

Marker for
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

bol is seen both in the cytoplasm and in a perinucleuar dot in spermatocytes, and in the cytoplasm of spermatids.

Marker for
Subcellular Localization
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
located_in nucleus
inferred from direct assay
located_in perikaryon
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\bol 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
bol
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
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 ( 38 )
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 bol
Transgenic constructs containing regulatory region of bol
Aberrations (Deficiencies and Duplications) ( 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
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (66)
6 of 14
No
Yes
3  
2 of 14
No
No
2  
1 of 14
No
No
1  
1 of 14
No
No
3  
1 of 14
No
No
2  
1 of 14
No
No
3  
1 of 14
No
No
2  
1 of 14
No
No
1  
1 of 14
No
No
1  
1 of 14
No
No
1  
1 of 14
No
No
1  
1 of 14
No
No
2  
1 of 14
No
No
1  
1 of 14
No
No
8  
1 of 14
No
No
1  
1 of 14
No
No
1  
1 of 14
No
No
1  
1 of 14
No
No
1  
1 of 14
No
No
1  
1 of 14
No
No
3  
1 of 14
No
No
1  
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (34)
11 of 14
Yes
Yes
Mus musculus (laboratory mouse) (38)
11 of 14
Yes
Yes
1 of 14
No
No
0  
Xenopus tropicalis (Western clawed frog) (28)
7 of 13
Yes
Yes
Danio rerio (Zebrafish) (49)
7 of 14
Yes
Yes
Caenorhabditis elegans (Nematode, roundworm) (18)
10 of 14
Yes
Yes
Anopheles gambiae (African malaria mosquito) (19)
Arabidopsis thaliana (thale-cress) (73)
Saccharomyces cerevisiae (Brewer's yeast) (6)
Schizosaccharomyces pombe (Fission yeast) (6)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:bol. Refer to their site for version information.
Paralogs
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?
5 of 14
5 of 14
5 of 14
5 of 14
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.
Dmel gene
Ortholog showing functional complementation
Supporting References
Interactions
Summary of Physical Interactions
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 with the translational regulator orb2.
(UniProt, Q24207 )
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
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3L
Recombination map
3-28
Cytogenetic map
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
66F5-67A1
Limits computationally determined from genome sequence between P{PZ}l(3)0162901629&P{PZ}mRpL1210534 and P{EP}Hsp26EP3336&P{EP}Hsp26EP3315
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
66F1-66F6
(determined by in situ hybridisation)
Experimentally Determined Recombination Data
Location
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (18)
Genomic Clones (24)
cDNA Clones (82)

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 essential function of bol at the transition from G2 to M phase during meiosis is in the control of twe translation.

Meiotic entry failure in bol mutants can be overcome by heterologous expression of twe.

Rescue experiments suggest Xlae\dazl may play a role analogous to bol in promoting meiotic cell division.

bol has an essential meiotic function in spermatogenesis.

Meiotic entry defect.

Relationship to Other Genes
Source for database merge of

Source for merge of: bol BcDNA:AT13332

Source for merge of: bol CG4727

Additional comments

Annotations CG4760 and CG4727 merged as CG4760 in release 3 of the genome annotation.

Source for merge of bol BcDNA:AT13332 was shared cDNA (AT13332) ( date:030602 ).

Nomenclature History
Source for database identify of
Nomenclature comments
Etymology
Synonyms and Secondary IDs (10)
Reported As
Symbol Synonym
BcDNA:AT13332
bol
(Collins et al., 2024, Titlow et al., 2023, Zheng et al., 2023, Ibaraki et al., 2021, Lawlor et al., 2021, Lécureuil et al., 2021, Pang et al., 2021, Kurmangaliyev et al., 2020, Mazina et al., 2020, Shi et al., 2020, Fingerhut et al., 2019, Shih et al., 2019, Croset et al., 2018, Swevers et al., 2018, Hu et al., 2017.6.13, Ota et al., 2017, Carbone et al., 2016, Clandinin and Owens, 2016-, Williams et al., 2016, Brooks et al., 2015, Gene Disruption Project members, 2015-, Van Bortle et al., 2015, Ashwal-Fluss et al., 2014, Rodriguez et al., 2012, Smibert et al., 2012, Xu et al., 2012, Carreira et al., 2011, Hartmann et al., 2011, Hilgers et al., 2011, Moon et al., 2011, Gan et al., 2010, Kwon et al., 2009, Rautureau et al., 2009, Hoopfer et al., 2008, Turner et al., 2008, Chintapalli et al., 2007, Christensen et al., 2007.10.29, Grieder et al., 2007, Sofola et al., 2007, Arya et al., 2006, Gresens et al., 2006.7.11, Mukai et al., 2006, Joiner and Wu, 2004, Xu et al., 2003)
ms(3)07272
Secondary FlyBase IDs
  • FBgn0040826
  • FBgn0063736
Datasets (0)
Study focus (0)
Experimental Role
Project
Project Type
Title
Study result (0)
Result
Result Type
Title
External Crossreferences and Linkouts ( 89 )
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 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.
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
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
References (158)