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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
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]
Also Known As
ERCC3, XPB, TFIIH, DmXPB, nc2
Key Links
Genomic Location
Cytogenetic map
Sequence location
3L:10,631,079..10,633,969 [+]
Recombination map
3-33
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
Belongs to the helicase family. RAD25/XPB subfamily. (Q02870)
Summaries
Gene Group (FlyBase)
TRANSCRIPTION FACTOR II H - CORE COMPLEX -
The general transcription factor complex Transcription factor II H (TFIIH) is required in RNA pol II transcription at initiation, promoter escape, early elongation steps and transcription re-initiation after polymerase pausing. TFIIH can be divided into two subcomplexes: core and cyclin-dependent protein kinase-activating kinase (CAK). The core complex contains an ATP-dependent DNA helicase activity required for promoter DNA opening and nucleotide excision repair (NER). (Adapted from PMID:25693126 and PMID:21592869).
Protein Function (UniProtKB)
ATP-dependent 3'-5' DNA helicase, component of the general transcription and DNA repair factor IIH (TFIIH) core complex, which is involved in general and transcription-coupled nucleotide excision repair (NER) of damaged DNA and, when complexed to CAK, in RNA transcription by RNA polymerase II. 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 ATPase activity of haywire/XPB/ERCC3, but not its helicase activity, is required for DNA opening. In transcription, TFIIH has an essential role in transcription initiation. When the pre-initiation complex (PIC) has been established, TFIIH is required for promoter opening and promoter escape. The ATP-dependent helicase activity of haywire/XPB/ERCC3 is required for promoter opening and promoter escape. Phosphorylation of the C-terminal tail (CTD) of the largest subunit of RNA polymerase II by the kinase module CAK controls the initiation of transcription.
(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 GBrowse view of Dmel\hay or 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
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
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
Gene Ontology (23 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 sequence or structural similarity with HGNC:3435
inferred from biological aspect of ancestor with PANTHER:PTN000138598
(assigned by GO_Central )
inferred from electronic annotation with InterPro:IPR006935
(assigned by InterPro )
inferred from electronic annotation with InterPro:IPR006935
(assigned by InterPro )
inferred from sequence or structural similarity with HGNC:3435
inferred from biological aspect of ancestor with PANTHER:PTN000138598
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000138598
(assigned by GO_Central )
Biological Process (12 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (7 terms)
CV Term
Evidence
References
inferred from sequence or structural similarity with HGNC:3435
inferred from biological aspect of ancestor with PANTHER:PTN000138598
(assigned by GO_Central )
inferred from sequence or structural similarity with HGNC:3435
inferred from biological aspect of ancestor with PANTHER:PTN000138598
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000138598
(assigned by GO_Central )
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
(assigned by GO_Central )
Cellular Component (5 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000138598
(assigned by GO_Central )
inferred by curator from GO:0000381
inferred from biological aspect of ancestor with PANTHER:PTN000138598
(assigned by GO_Central )
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
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000138598
(assigned by GO_Central )
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
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

GBrowse - Visual display of RNA-Seq signals

View Dmel\hay in GBrowse 2
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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
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, and Transgenic Constructs
Classical and Insertion Alleles ( 19 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 14 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of hay
Transgenic constructs containing regulatory region of hay
Deletions and Duplications ( 18 )
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 v7.1)
Homo sapiens (Human) (1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
15 of 15
Yes
Yes
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
14 of 15
Yes
Yes
Rattus norvegicus (Norway rat) (1)
13 of 13
Yes
Yes
Xenopus tropicalis (Western clawed frog) (1)
7 of 12
Yes
Yes
Danio rerio (Zebrafish) (1)
12 of 15
Yes
Yes
Caenorhabditis elegans (Nematode, roundworm) (1)
15 of 15
Yes
Yes
Arabidopsis thaliana (thale-cress) (2)
9 of 9
Yes
Yes
9 of 9
Yes
Yes
Saccharomyces cerevisiae (Brewer's yeast) (1)
15 of 15
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (1)
12 of 12
Yes
Yes
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091902X7 )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091501M8 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W01NS )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X01LU )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0291 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (0)
No records found.
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 3 )
Potential Models Based on Orthology ( 1 )
Modifiers Based on Experimental Evidence ( 0 )
Allele
Disease
Interaction
References
Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
 
Disease Associations of Human Orthologs (via DIOPT v7.1 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
esyN Network Key:
Suppression
Enhancement

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.
InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Gene Group - Pathway Membership (FlyBase)
External Data
Linkouts
KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3L
Recombination map
3-33
Cytogenetic map
Sequence location
3L:10,631,079..10,633,969 [+]
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 (8)
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 GBrowse for alignment of the cDNAs and ESTs to the gene model.

cDNA clones, fully sequences
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
GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
Antibody Information
Laboratory Generated Antibodies
 
Commercially Available Antibodies
 
Other Information
Relationship to Other Genes
Source for database identify of
Source for identity of: hay CG8019
Source for database merge of
Additional comments
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.
Origin and Etymology
Discoverer
Etymology
Identification
External Crossreferences and Linkouts ( 80 )
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/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
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
KEGG Genes - Molecular building blocks of life in the genomic space.
KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
modMine - A data warehouse for the modENCODE project
Linkouts
BioGRID - A database of protein and genetic interactions.
DroID - A comprehensive database of gene and protein interactions.
DRSC - Results frm RNAi screens
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyMine - An integrated database for Drosophila genomics
InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Synonyms and Secondary IDs (17)
Reported As
Symbol Synonym
Secondary FlyBase IDs
  • FBgn0004376
  • FBgn0015830
Datasets (0)
Study focus (0)
Experimental Role
Project
Project Type
Title
References (111)