FB2026_03 , released September 17, 2026
Gene: Dmel\Dnaaf5
Open Close
General Information
Symbol
Dmel\Dnaaf5
Species
D. melanogaster
Name
Dynein axonemal assembly factor 5
Annotation Symbol
CG31320
Feature Type
protein_coding_gene
FlyBase ID
FBgn0288976
Gene Model Status
Current
Stock Availability
8 publicly available
Gene Summary
Cytoplasmic protein involved in the delivery of the dynein machinery to the motile cilium. It is required for the assembly of the axonemal dynein inner and outer arms, two structures attached to the peripheral outer doublet A microtubule of the axoneme, that play a crucial role in cilium motility. (UniProt, Q8INF7)
Contribute a Gene Snapshot for this gene.
All Summaries
Also Known As

HEATR2, fried

Key Links
Genomic Location
Cytogenetic map
88B6-88B6
Sequence location
Recombination map
3-55
RefSeq locus
NT_033777 REGION:14358356..14361786
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 (11 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
enables dynein intermediate chain binding
inferred from biological aspect of ancestor with PANTHER:PTN000416608
(GO Reference Genome Project, 2011-)
inferred from sequence or structural similarity with UniProtKB:Q86Y56
(Diggle et al., 2014)
Biological Process (7 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
involved_in adult locomotory behavior
inferred from mutant phenotype
(Diggle et al., 2014)
involved_in cilium movement involved in cell motility
inferred from mutant phenotype
(Diggle et al., 2014)
involved_in flagellated sperm motility
inferred from mutant phenotype
(Diggle et al., 2014)
involved_in inner dynein arm assembly
inferred from mutant phenotype
(Diggle et al., 2014)
involved_in outer dynein arm assembly
inferred from mutant phenotype
(Diggle et al., 2014)
involved_in sensory perception of sound
inferred from mutant phenotype
(Diggle et al., 2014)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
involved_in cilium movement
inferred from sequence or structural similarity with UniProtKB:Q86Y56
(Gene Ontology Curators, 2002-)
involved_in flagellated sperm motility
inferred from biological aspect of ancestor with PANTHER:PTN000416608
(GO Reference Genome Project, 2011-)
involved_in inner dynein arm assembly
inferred from biological aspect of ancestor with PANTHER:PTN000416608
(GO Reference Genome Project, 2011-)
involved_in outer dynein arm assembly
inferred from biological aspect of ancestor with PANTHER:PTN000416608
(GO Reference Genome Project, 2011-)
Cellular Component (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
(Diggle et al., 2014)
NOT located_in motile cilium
inferred from direct assay
(Diggle et al., 2014)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in dynein axonemal particle
inferred from biological aspect of ancestor with PANTHER:PTN000416608
(GO Reference Genome Project, 2011-)
located_in dynein axonemal particle
inferred from sequence or structural similarity with UniProtKB:A0JMW2
(Gene Ontology Curators, 2002-)
Gene Group / Complex (FlyBase)
Protein Family (UniProt)
Belongs to the DNAAF5 family. (Q8INF7)
Protein Signatures (InterPro)
Summaries
Protein Function (UniProtKB)
Cytoplasmic protein involved in the delivery of the dynein machinery to the motile cilium. It is required for the assembly of the axonemal dynein inner and outer arms, two structures attached to the peripheral outer doublet A microtubule of the axoneme, that play a crucial role in cilium motility.
(UniProt, Q8INF7)
Summary (Interactive Fly)

Dynein assembly factor - involved in assembly of motile cilia - component of intracellular pre-assembly and transport network - necessary to deliver functional dynein machinery to the ciliary compartment - mutant flies exhibit aberrant proprioception, deafness and immotile sperm

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

Please see the JBrowse view of Dmel\Dnaaf5 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 Q8INF7)

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 5.48

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Dnaaf5-RA
FBtr0082943
NM_169581
3431
845
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
Dnaaf5-PA
FBpp0082402
96.3
845
5.28
Q8INF7
NP_731905
AAN13608
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)
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\Dnaaf5 using the Feature Mapper tool.

External Data
Crossreferences
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).

2.22

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
organism | subset
(Fisher et al., 2012)
embryonic stage 12
mesoderm
(Diggle et al., 2014)
embryonic stage 12 -- 16
chordotonal organ
(Diggle et al., 2014)
embryonic stage 13 -- 16
dorsal sensory complex proper primordium

Comment: reported as dorsal/lateral sensory complexes

(Fisher et al., 2012)
embryonic head sensory system
(Fisher et al., 2012)
sensory nervous system primordium
(Fisher et al., 2012)
ventral sensory complex primordium
(Fisher et al., 2012)
embryonic stage 14 -- 16
chordotonal neuron
(Zur Lage et al., 2019)
embryonic stage 15
chordotonal organ precursor cell
(Newton et al., 2012)
embryonic stage 16
mechanosensory neuron
(Diggle et al., 2014)
mutational analysis
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
chordotonal neuron
(Zur Lage et al., 2019)
adult stage | male
spermatogonium

Comment: data from FlyAtlas

(Zur Lage et al., 2019)
Additional Descriptive Data

As part of a survey of ciliary motility gene homologs in Drosophila, expression of HEATR2 was assayed in motile ciliated cell types. Expression was observed in both chordotonal neurons and testis (sperm).

(Zur Lage et al., 2019)

HEATR2 transcripts are detected in chordotonal neurons and transiently in mesoderm from embryonic stage 12.

(Diggle et al., 2014)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult head
(Aradska et al., 2015)
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
(Diggle et al., 2014)
NOT located_in motile cilium
inferred from direct assay
(Diggle et al., 2014)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Dnaaf5 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
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
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 6 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 5 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Dnaaf5
Transgenic constructs containing regulatory region of Dnaaf5
Aberrations (Deficiencies and Duplications) ( 2 )
Inferred from experimentation ( 2 )
Gene disrupted in
Df(3R)Exel6267
(Ryder, 2004.4.26)
Gene partially disrupted in
Df(3R)Dnaaf527
(Diggle et al., 2014)
Inferred from location ( 4 )
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 | recessive
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4sca-DB
viable, with Scer\GAL4tin.CΔ4
Sterility
Allele
female sterile | germline clone | recessive
male sterile, with Scer\GAL4VP16.bam
Other Phenotypes
Allele
abnormal auditory perception | larval stage, with Scer\GAL4sca-DB
abnormal body size | larval stage
abnormal locomotor behavior | adult stage, with Scer\GAL4sca.PU
short lived | heat sensitive, with Scer\GAL4tin.CΔ4
uncoordinated | adult stage, with Scer\GAL4sca-DB
visible, with Scer\GAL4pnr-MD237
Phenotype manifest in
Allele
egg chamber | germline clone
inner dynein arm, with Scer\GAL4sca-DB
inner dynein arm, with Scer\GAL4VP16.bam
Johnston organ neuron, with Scer\GAL4sca-DB
karyosome | germline clone
nurse cell & nucleus | germ-line clone
outer dynein arm, with Scer\GAL4sca-DB
outer dynein arm, with Scer\GAL4VP16.bam
spermatid, with Scer\GAL4VP16.bam
spermatozoon, with Scer\GAL4VP16.bam
trichogen cell, with Scer\GAL4pnr-MD237
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) (1)
Hsap\DNAAF5
14 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) (1)
Rnor\Dnaaf5
14 of 14
Yes
Yes
Mus musculus (laboratory mouse) (1)
Mmus\Dnaaf5
12 of 14
Yes
Yes
Xenopus tropicalis (Western clawed frog) (1)
Xtro\dnaaf5
11 of 13
Yes
Yes
Danio rerio (Zebrafish) (3)
Drer\dnaaf5
2 of 14
Yes
Yes
Drer\dnaaf5
1 of 14
No
Yes
Drer\gcn1
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (1)
Cele\gcn-1
1 of 14
Yes
No
Anopheles gambiae (African malaria mosquito) (2)
Agam\AgaP_AGAP002100
12 of 12
Yes
Yes
Agam\AgaP_AGAP001700
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (1)
Atha\ILA
1 of 13
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (1)
Scer\TPD3
1 of 13
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Dnaaf5. 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) (1)
l(3)80Fj
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 2 )
Potential Models Based on Orthology ( 1 )
Human Ortholog
Disease
Evidence
References
DNAAF5; dynein axonemal assembly factor 5
model of  primary ciliary dyskinesia 18
IEA
(FlyBase, 2019-)
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?
DNAAF5; dynein axonemal assembly factor 5
14 of 14
614864
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
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
Pathways
Signaling Pathways (FlyBase)
Metabolic Pathways
FlyBase
External Links
External Data
Linkouts
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3R
Recombination map
3-55
Cytogenetic map
88B6-88B6
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
88B6-88B6
Limits computationally determined from genome sequence between P{lacW}trxj14A6&P{PZ}trx00347 and P{PZ}cv-c06951
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
Mapping based on deficiency analysis (details unspecified).
(Morris et al., 2003)
88B-88C3
(determined by in situ hybridisation)
(Morris et al., 2003)
Experimentally Determined Recombination Data
Location

3-55

(FlyBase, 2016)

3-53.1

(Comeron et al., 2012)
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (8)
Genomic Clones (23)
cDNA Clones (9)
 

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
      Antibody Information
      Laboratory Generated Antibodies
       
      Commercially Available Antibodies
       
      Cell Line Information
      Publicly Available Cell Lines
       
        Other Stable Cell Lines
         
          Other Comments

          New annotation (CG31320) 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: CG31320 NEST:bs27f03

          (Rana, 2004.7.8)

          Source for merge of: HEATR2 fried

          (Morris, 2022.8.30)
          Additional comments
          Nomenclature History
          Source for database identify of

          Source for identity of: HEATR2 CG31320

          (Diggle et al., 2014)
          Nomenclature comments
          Etymology

          Named 'HEATR2' after the human ortholog.

          (Diggle et al., 2014)

          The gene is named "fried" after one of "the unfortunate fates commonly met by eggs", since mutants show defects in oocyte specification or early oogenesis.

          (Morris et al., 2003)
          Synonyms and Secondary IDs (8)
          Reported As
          Symbol Synonym
          CG31320
          (Morris, 2022.8.30, Zur Lage et al., 2019, Hosono et al., 2015, Diggle et al., 2014, Kwon et al., 2013, Newton et al., 2012, Frise et al., 2010, Keleman et al., 2009.8.5, Perkins et al., 2009)
          Dnaaf5
          HEATR2
          (Blümel et al., 2022, Clandinin and Owens, 2016-, Diggle et al., 2014)
          Heatr2
          (Zur Lage et al., 2019)
          NEST:bs27f03
          fried
          (Morris, 2022.8.30)
          Name Synonyms
          Dynein axonemal assembly factor 5
          HEAT repeat containing 2
          fried
          (Morris et al., 2003)
          Secondary FlyBase IDs
          • FBgn0051320
          • FBgn0045167
          • FBgn0067771
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 26 )
          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
          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
          FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
          Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
          iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
          Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
          References (38)