FB2024_04 , released June 25, 2024
Gene: Dmel\hay
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General Information
Symbol
Dmel\hay
Species
D. melanogaster
Name
haywire
Annotation Symbol
CG8019
Feature Type
FlyBase ID
FBgn0001179
Gene Model Status
Stock Availability
Enzyme Name (EC)
DNA 3'-5' helicase (5.6.2.4)
Gene Summary
haywire (hay) encodes a ATP-dependent 3'-5' DNA helicase based on the similarity with the human gene XPB, which is a subunit of the DNA repair and basal transcription factor TFIIH. The product of hay contributes to response to UV and homeotic gene expression. [Date last reviewed: 2018-09-13] (FlyBase Gene Snapshot)
Also Known As

XPB, ERCC3, TFIIH, DmXPB, nc2

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
3-33
RefSeq locus
NT_037436 REGION:10631079..10633969
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (16 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (6 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000138598
inferred from sequence or structural similarity with HGNC:3435
enables ATP binding
inferred from electronic annotation with InterPro:IPR006935
enables DNA binding
inferred from electronic annotation with InterPro:IPR006935
inferred from electronic annotation with InterPro:IPR001161
inferred from sequence or structural similarity with HGNC:3435
inferred from electronic annotation with InterPro:IPR006935
Biological Process (6 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
involved_in response to UV
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
inferred from sequence or structural similarity with HGNC:3435
inferred from electronic annotation with InterPro:IPR001161
inferred from sequence or structural similarity with HGNC:3435
inferred from sequence or structural similarity with HGNC:3435
inferred from sequence or structural similarity with SGD:S000001405
inferred from biological aspect of ancestor with PANTHER:PTN000138598
inferred from electronic annotation with InterPro:IPR001161
inferred from sequence or structural similarity with HGNC:3435
Cellular Component (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
inferred from sequence or structural similarity with HGNC:3435
inferred from sequence or structural similarity with SGD:S000001405
inferred from biological aspect of ancestor with PANTHER:PTN000138598
Protein Family (UniProt)
Belongs to the helicase family. RAD25/XPB subfamily. (Q02870)
Catalytic Activity (EC/Rhea)
3'-5' DNA helicase activity
Couples ATP hydrolysis with the unwinding of duplex DNA by translocating in the 3'-5' direction (5.6.2.4)
Summaries
Gene Snapshot
haywire (hay) encodes a ATP-dependent 3'-5' DNA helicase based on the similarity with the human gene XPB, which is a subunit of the DNA repair and basal transcription factor TFIIH. The product of hay contributes to response to UV and homeotic gene expression. [Date last reviewed: 2018-09-13]
Gene Group (FlyBase)
TRANSCRIPTION FACTOR II H - CORE COMPLEX -
The general transcription factor complex Transcription factor II H (TFIIH) core complex, is a subcomplex of the TFIIH holoenzyme required in RNA pol II transcription with an ATP-dependent DNA helicase activity required for promoter DNA opening. TFIIH also functions in nucleotide excision repair (NER). After complex loading and the verification step of NER, the cyclin-dependent protein kinase-activating kinase (CAK) subcomplex dissociates and the TFIIH core complex functions independently. (Adapted from PMID:25693126, PMID:21592869 and PMID:30798933).
UNCLASSIFIED DNA HELICASES -
This group comprises DNA helicases that do not classify under other groups in FlyBase.
Protein Function (UniProtKB)
ATP-dependent 3'-5' DNA helicase/translocase; binds dsDNA rather than ssDNA, unzipping it in a translocase rather than classical helicase activity. Component of the general transcription and DNA repair factor IIH (TFIIH) core complex. When complexed to CDK-activating kinase (CAK), involved in RNA transcription by RNA polymerase II. The ATPase activity of XPB/ERCC3, but not its helicase activity, is required for DNA opening; it may wrap around the damaged DNA wedging it open, causing localized melting and twisting that allows XPD/ERCC2 helicase to anchor. The ATP-dependent helicase activity of XPB/ERCC3 may be required for promoter escape. Also involved in transcription-coupled nucleotide excision repair (NER) of damaged DNA. In NER, TFIIH acts by opening DNA around the lesion to allow the excision of the damaged oligonucleotide and its replacement by a new DNA fragment. The structure of the TFIIH transcription complex differs from the NER-TFIIH complex; large movements by XPD/ERCC2 and XPB/ERCC3 are stabilized by XPA.
(UniProt, Q02870)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
hay: haywire
The original allele, haync2, is homozygous male sterile; some derivative alleles are recessive lethal. Homozygous haync2 males display defects in meiosis, flagellar elongation, and nuclear shaping. haync2 also male sterile in trans heterozygotes with recessive male-sterile alleles of β Tub85D or with wrlnc4 suggesting a role for the haywire product in microtubule function. Deficiencies of hay are fertile in heterozygous combination with the above interacting mutations, indicating that the extragenic failure haync2 to complement is based on a poison product mechanism. This feature was used to select alleles (haync2rv1-8) that revert the failure of haync2 to complement β2tn (= βTub85Dn).
Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\hay 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 Q02870)

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 5.43

Shares 5' UTR with downstream gene.

Gene model reviewed during 5.46

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0076253
2604
798
FBtr0114599
2511
798
Additional Transcript Data and Comments
Reported size (kB)

2.7 (unknown)

2.5 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0075982
90.3
798
7.94
FBpp0113091
90.3
798
7.94
Polypeptides with Identical Sequences

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

798 aa isoforms: hay-PA, hay-PB
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

Component of the 7-subunit TFIIH core complex composed of haywire/XPB/ERCC3, XPD/ERCC2, GTF2H1, GTF2H2, GTF2H3, GTF2H4 and GTF2H5, which is active in NER. The core complex associates with the 3-subunit CDK-activating kinase (CAK) module composed of CCNH/cyclin H, CDK7 and MNAT1 to form the 10-subunit holoenzyme (holo-TFIIH) active in transcription. Interacts with PUF60. Interacts with ATF7IP. Interacts with Epstein-Barr virus EBNA2.

(UniProt, Q02870)
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\hay 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).

-0.13

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

Comment: maternally deposited

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

hay transcripts are detected throughout development on northern blots.

hay transcripts are detected at all stages of development and are present at lower levels in embryos than in later stages.

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
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\hay 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
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 9-10
  • Stages(s) 13-16
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 hay
Transgenic constructs containing regulatory region of hay
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
adult abdomen & endocuticle
adult abdomen & endocuticle (with Df(3L)lxd6)
adult abdomen & endocuticle (with haync2)
adult abdomen & endocuticle (with haync2rv1)
adult abdomen & endocuticle (with haync2rv2)
adult abdomen & endocuticle (with haync2rv4)
adult abdomen & endocuticle (with haync2rv7)
adult abdomen & endocuticle (with haync2rv8)
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (1)
14 of 14
Yes
Yes
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (1)
14 of 14
Yes
Yes
Mus musculus (laboratory mouse) (1)
13 of 14
Yes
Yes
Xenopus tropicalis (Western clawed frog) (2)
11 of 13
Yes
Yes
5 of 13
No
Yes
Danio rerio (Zebrafish) (1)
13 of 14
Yes
Yes
Caenorhabditis elegans (Nematode, roundworm) (1)
14 of 14
Yes
Yes
Anopheles gambiae (African malaria mosquito) (1)
11 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (2)
13 of 13
Yes
Yes
13 of 13
Yes
Yes
Saccharomyces cerevisiae (Brewer's yeast) (3)
13 of 13
Yes
Yes
1 of 13
No
No
1 of 13
No
Yes
Schizosaccharomyces pombe (Fission yeast) (2)
12 of 12
Yes
Yes
2 of 12
No
Yes
Escherichia coli (enterobacterium) (1)
2 of 11
Yes
Yes
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:hay. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 3 )
Potential Models Based on Orthology ( 2 )
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
Summary of Genetic Interactions
esyN Network Diagram
Show/hide secondary interactors 
(data from AllianceMine provided by esyN)
esyN Network Key:
Suppression
Enhancement
Other 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)
Component of the 7-subunit TFIIH core complex composed of haywire/XPB/ERCC3, XPD/ERCC2, GTF2H1, GTF2H2, GTF2H3, GTF2H4 and GTF2H5, which is active in NER. The core complex associates with the 3-subunit CDK-activating kinase (CAK) module composed of CCNH/cyclin H, CDK7 and MNAT1 to form the 10-subunit holoenzyme (holo-TFIIH) active in transcription. Interacts with PUF60. Interacts with ATF7IP. Interacts with Epstein-Barr virus EBNA2.
(UniProt, Q02870 )
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
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3L
Recombination map
3-33
Cytogenetic map
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
67E5-67E5
Limits computationally determined from genome sequence between P{lacW}l(3)L0539L0539&P{PZ}Dhh1rL562 and P{PZ}simj01814
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
63E3-63E7
67E3-67E4
(determined by in situ hybridisation)
Experimentally Determined Recombination Data
Location

3-24.4 +/- 0.5

3-34.4

Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (6)
Genomic Clones (19)
cDNA Clones (36)
 

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)
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

      Mutant lesions in lethal hay alleles reveal the importance of the nucleotide-binding domain, suggesting an essential role for ATPase activity.

      Gene is involved in pre-replication DNA repair of UV and AA lesions.

      hay encodes the largest subunit of TfIIH.

      mRNA levels do not increase at adult day 5 in strain showing extended longevity phenotype (ELP).

      Low-stringency hybridization isolated the homolog of the human excision repair gene ERCC3, which maps to 67E3--67E4, known to be the previously defined haywire gene. Microinjection and DNA transfection of haywire into human xeroderma pigmentosum fibroblasts and group 3 rodent mutants did not yield detectable correction.

      The hay gene was cloned and is related to the human ERCC3 gene which is associated with both xeroderma pigmentosum and Cockayne's syndrome. Phenotypes of hay mutants mimic some of the effects of xeroderma pigmentosum in humans.

      The original allele, haync2, is homozygous male sterile; some derivative alleles are recessive lethal. Homozygous haync2 males display defects in meiosis, flagellar elongation and nuclear shaping. haync2 also male sterile in transheterozygotes with recessive male-sterile alleles of β Tub85D or with wrl1 suggesting a role for the haywire product in microtubule function. Deficiencies of hay are fertile in heterozygous combination with the above interacting mutations, indicating that the extragenic failure haync2 to complement is based on a poison product mechanism. This feature was used to select alleles (haync2rv1-8) that revert the failure of haync2 to complement βTub85Dn.

      Relationship to Other Genes
      Source for database merge of
      Additional comments
      Nomenclature History
      Source for database identify of

      Source for identity of: hay CG8019

      Nomenclature comments
      Etymology
      Synonyms and Secondary IDs (17)
      Reported As
      Symbol Synonym
      ms(3)nc2
      Secondary FlyBase IDs
      • FBgn0004376
      • FBgn0015830
      Datasets (0)
      Study focus (0)
      Experimental Role
      Project
      Project Type
      Title
      Study result (0)
      Result
      Result Type
      Title
      External Crossreferences and Linkouts ( 68 )
      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)
      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
      FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
      iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
      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
      References (117)