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General Information
Symbol
Dmel\H
Species
D. melanogaster
Name
Hairless
Annotation Symbol
CG5460
Feature Type
FlyBase ID
FBgn0001169
Gene Model Status
Stock Availability
Gene Snapshot
Hairless (H) encodes the major antagonist of Notch signaling activity during imaginal development. It silences target genes of the product of N by assembling a transcriptional repressor complex, which includes the transcription factor encoded by Su(H) and general co-repressors like those encoded by gro and CtBP. [Date last reviewed: 2019-03-07]
Key Links
Genomic Location
Cytogenetic map
Sequence location
3R:20,621,938..20,628,915 [+]
Recombination map

3-69

RefSeq locus
NT_033777 REGION:20621938..20628915
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (12 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:asf1; FB:FBgn0029094
inferred from direct assay
inferred from genetic interaction with FLYBASE:Su(H); FB:FBgn0004837
Terms Based on Predictions or Assertions (0 terms)
Biological Process (9 terms)
Terms Based on Experimental Evidence (9 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
    -
    Summaries
    Pathway (FlyBase)
    Negative Regulators of Notch Signaling Pathway -
    The Notch receptor signaling pathway is activated by the binding of the transmembrane receptor Notch (N) to transmembrane ligands, Dl or Ser, presented on adjacent cells. This results in the proteolytic cleavage of N, releasing the intracellular domain (NICD). NICD translocates into the nucleus, interacting with Su(H) and mam to form a transcription complex, which up-regulates transcription of Notch-responsive genes. Negative regulators of the pathway down-regulate the signal from the sending cell or the response in the receiving cell. (Adapted from FBrf0225731 and FBrf0192604).
    Protein Function (UniProtKB)
    Is a potent antagonist of neurogenic gene activity during sensory organ development. The expression of distinct cell fates by the trichogen (shaft) / tormogen (socket) sister cell pair depends on the level of H activity. A certain threshold level of H activity is required, below which both sister cells adopt the tormogen fate.
    (UniProt, Q02308)
    Phenotypic Description (Red Book; Lindsley and Zimm 1992)
    H: Hairless
    thumb
    H: Hairless
    From Bridges and Morgan, 1923, Carnegie Inst. Washington Publ. No. 327: 161.
    Bristles, especially postverticals and abdominals, missing in heterozygous H flies. Bristle sockets present at some sites, not at others. Expressed most distinctly on head; occipital; post vertical and ocellar bristles affected. Bristles of antennae and vibrissae show mutant phenotype much less frequently. Sockets without shafts also found on thorax, scutellum, abdominal tergites, external genitalia, wings, and legs. No shaftless sockets appear on the bracted costa of the wing. Some 40% of bristle organs located on distal part of femur differentiate neither shaft nor bract; bracts absent whenever shaft missing but present when shaft present; abnormally short shaft may be accompanied by normal-sized bract (from description of H2, by Tobler, Rothenbuhler, and Nothiger, 1973, Experientia 29: 370-71). Veins L4 and L5 do not reach wing margin; occasionally true of L2 also. Eyes larger than wild type; body color somewhat paler. Lees and Waddington [1942, Proc. Roy. Soc. (London), Ser. B. 131: 87-110 (fig.)] show that trichogen cell forms a socket instead of a bristle shaft at some sites. Phenotypic expression of H responds linearly to dosage of Su(H)+ in region 35B6-10 on the left arm of chromosome 2. The number of microchaetae in H/+ flies varies from approximately 35 in the presence of a single dose of 35B6-10 to fewer than 10 in the presence of four doses (Ashburner, 1982, Genetics 64: 471-79). Interactions with other mutants studied by House (1953, Genetics 38: 199-215, 309-27; 1959, Genetics 44: 516; 1955, Anat. Record 122: 471; 1959, Anat. Record 134: 581-82). H suppresses wing notching of N, fa, fano, and nd, enhances Ax; also H enhances eye effect of spl, and removes more bristles in combination with spl (House, Von Halle). Reduction in the number of copies of the wild-type allele of H decreases the mutant phenotype of heterozygous N and Dl flies, but increase in the number of copies of the wild-type allele of H enhances the mutant phenotype of heterozygous N, Dl, and E(spl) flies (Vassin, Vielmetter, and Campos-Ortega, 1985, J. Neurogenet. 2: 291-308). H shows some superadditive interaction with en, ci, ciW, and ciD relative to degree of L4 interruption. L2 interruption augmented in combinations with ve and ri; L3 interruption augmented in combinations with ve and tt. Triploid, H/+/+, intermediate between wild type and H/+. H/H/+ most extreme type with bristles absent from head, thorax, and abdomen [Gowen, 1933, Am. Nat. 67: 178-80 (fig.)]. Homozygous lethal. H null homozygotes die during larval and pupal stages (Bang et al.). Animals surviving to pharate adult are completely devoid of macrochaetes and microchaetes on the head and notum, with occasional "double sockets" remaining on the abdominal tergites. Bristles on the legs significantly resistant to loss of H+ function; many "double sockets" and some normal bristles remain on the legs of H null homozygotes. In regions of the notum exhibiting bristle "loss" in adult H mutants, macrochaete and microchaete primary precursor cells undetectable (Bang et al.). RK1.
    H16
    Recessive allele of Hairy (formerly called Hairless-recessive). Homozygotes have nearly all bristles and hairs substituted by double and triple abnormal sockets; veins L4 and L5 fail to reach margin. H16/H lethal, probably in pupal stage; few escapers short lived with extreme Hairless phenotype; all bristles and hairs suppressed or substituted by abnormal sockets; wings reduced; veins L2, L4 and L5 abnormal.
    H22
    Less severe than H16. Almost completely recessive. Heterozygotes most frequently wildtype in phenotype, but occasionally a "double socket" appears on the head in the position of a postvertical macrochaete or on the abdominal tergites. Homozygotes display a much stronger and more extensive mutant phenotype than H null heterozygotes. Many head and notum macrochaetes and approximately 50% of notum microchaetes missing; remaining 50% exhibit a spectrum of "double socket" phenotypes. Homozygotes also exhibit loss of wing vein tissue from L4 and L5. Approximately 50% of flies carrying H22r in trans to a H null allele die as pharate adults. The remainder survive to eclosion, but are short-lived and exhibit extensive loss of both macrochaetes and microchaetes on the head, notum, and abdominal tergites; only 20% of the notum microchaetes remain, all with a completely transformed "double socket" phenotype.
    Summary (Interactive Fly)

    novel basic protein - a major antagonist of the Notch signaling-pathway - associates with Suppressor of Hairless, thereby inhibiting trans-activation of Notch target genes

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

    Please see the GBrowse view of Dmel\H or the JBrowse view of Dmel\H 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 5.47

    Sequence Ontology: Class of Gene
    Transcript Data
    Annotated Transcripts
    Name
    FlyBase ID
    RefSeq ID
    Length (nt)
    Assoc. CDS (aa)
    FBtr0083914
    5919
    1077
    FBtr0083915
    3884
    1059
    FBtr0083917
    6004
    1059
    FBtr0083916
    4255
    1077
    Additional Transcript Data and Comments
    Reported size (kB)

    6.0, 5.3, 5.0, 4.2, 4.0 (northern blot)

    6, 5.5, 4 (northern blot)

    Comments
    External Data
    Crossreferences
    Polypeptide Data
    Annotated Polypeptides
    Name
    FlyBase ID
    Predicted MW (kDa)
    Length (aa)
    Theoretical pI
    RefSeq ID
    GenBank
    FBpp0083322
    111.1
    1077
    10.08
    FBpp0083323
    109.2
    1059
    10.14
    FBpp0083325
    109.2
    1059
    10.14
    FBpp0083324
    111.1
    1077
    10.08
    Polypeptides with Identical Sequences

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

    1077 aa isoforms: H-PA, H-PD
    1059 aa isoforms: H-PB, H-PC
    Additional Polypeptide Data and Comments
    Reported size (kDa)

    1059 (aa); 109 (kD)

    1076 (aa); 110 (kD)

    Comments

    No extensive homology to other protein sequences in the databases was observed. The H protein is highly basic overall but has a >100 aa stretch that is very acidic surrounded by particularly basic regions. Long homopolymeric runs of alanine residues are also present,especially near the carboxy terminus.

    One of a couple of products generated by alternative splicing.

    External Data
    Crossreferences
    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\H 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
    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
    No Assay Recorded
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    in situ
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    northern blot
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    Additional Descriptive Data
    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

    H protein is found in all cells of all tissues at all stages assayed.

    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\H 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
    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
    Alleles, Insertions, and Transgenic Constructs
    Classical and Insertion Alleles ( 109 )
    For All Classical and Insertion Alleles Show
     
    Other relevant insertions
    Transgenic Constructs ( 62 )
    For All Alleles Carried on Transgenic Constructs Show
    Transgenic constructs containing/affecting coding region of H
    Transgenic constructs containing regulatory region of H
    Deletions and Duplications ( 12 )
    Phenotypes
    For more details about a specific phenotype click on the relevant allele symbol.
    Lethality
    Allele
    Sterility
    Allele
    Other Phenotypes
    Allele
    visible (with H1)
    visible (with H3)
    Phenotype manifest in
    Allele
    adult abdominal sensillum & sternite
    adult head & macrochaeta
    adult head & microchaeta
    adult thorax & macrochaeta
    adult thorax & macrochaeta, with Scer\GAL4T113
    dorsal mesothoracic disc & cell
    dorsal row & sensory mother cell, with Scer\GAL4ap-md544
    head & macrochaeta
    macrochaeta & adult head
    macrochaeta & head
    macrochaeta & scutum
    mesothoracic tergum & macrochaeta
    mesothoracic tergum & macrochaeta | somatic clone
    microchaeta & scutum
    microchaeta & tormogen cell | conditional ts
    microchaeta & trichogen cell
    scutum & macrochaeta
    wing & macrochaeta
    wing & microchaeta | ectopic, with Scer\GAL4en-e16E
    Orthologs
    Human Orthologs (via DIOPT v8.0)
    Homo sapiens (Human) (0)
    No records found.
    Model Organism Orthologs (via DIOPT v8.0)
    Mus musculus (laboratory mouse) (0)
    No records found.
    Rattus norvegicus (Norway rat) (0)
    No records found.
    Xenopus tropicalis (Western clawed frog) (0)
    No records found.
    Danio rerio (Zebrafish) (0)
    No records found.
    Caenorhabditis elegans (Nematode, roundworm) (0)
    No records found.
    Arabidopsis thaliana (thale-cress) (0)
    No records found.
    Saccharomyces cerevisiae (Brewer's yeast) (0)
    No records found.
    Schizosaccharomyces pombe (Fission yeast) (0)
    No records found.
    Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091906C5 )
    Organism
    Common Name
    Gene
    AAA Syntenic Ortholog
    Multiple Dmel Genes in this Orthologous Group
    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) ( EOG09150AZW )
    Organism
    Common Name
    Gene
    Multiple Dmel Genes in this Orthologous Group
    Musca domestica
    House fly
    Glossina morsitans
    Tsetse fly
    Glossina morsitans
    Tsetse fly
    Lucilia cuprina
    Australian sheep blowfly
    Mayetiola destructor
    Hessian fly
    Aedes aegypti
    Yellow fever mosquito
    Anopheles darlingi
    American malaria mosquito
    Anopheles gambiae
    Malaria mosquito
    Culex quinquefasciatus
    Southern house mosquito
    Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0FFE )
    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
    Cimex lectularius
    Bed bug
    Acyrthosiphon pisum
    Pea aphid
    Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0FF2 )
    Organism
    Common Name
    Gene
    Multiple Dmel Genes in this Orthologous Group
    Strigamia maritima
    European centipede
    Strigamia maritima
    European centipede
    Strigamia maritima
    European centipede
    Strigamia maritima
    European centipede
    Strigamia maritima
    European centipede
    Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
    No non-Arthropod Metazoa orthologies identified
    Paralogs
    Paralogs (via DIOPT v8.0)
    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 ( 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 v8.0 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?
      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
      esyN Network Diagram
      Show neighbor-neighbor interactions:
      Select Layout:
      Legend:
      Protein
      RNA
      Selected Interactor(s)
      Interactions Browser

      Please see the Physical Interaction reports below for full details
      protein-protein
      Physical Interaction
      Assay
      References
      RNA-RNA
      Physical Interaction
      Assay
      References
      RNA-protein
      Physical Interaction
      Assay
      References
      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
      suppressible
      enhanceable
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      suppressible
      suppressible
      suppressible
      suppressible
      suppressible
      suppressible
      suppressible
      External Data
      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
      Signaling Pathways (FlyBase)
      Negative Regulators of Notch Signaling Pathway -
      The Notch receptor signaling pathway is activated by the binding of the transmembrane receptor Notch (N) to transmembrane ligands, Dl or Ser, presented on adjacent cells. This results in the proteolytic cleavage of N, releasing the intracellular domain (NICD). NICD translocates into the nucleus, interacting with Su(H) and mam to form a transcription complex, which up-regulates transcription of Notch-responsive genes. Negative regulators of the pathway down-regulate the signal from the sending cell or the response in the receiving cell. (Adapted from FBrf0225731 and FBrf0192604).
      Metabolic Pathways
      External Data
      Linkouts
      KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
      Genomic Location and Detailed Mapping Data
      Chromosome (arm)
      3R
      Recombination map

      3-69

      Cytogenetic map
      Sequence location
      3R:20,621,938..20,628,915 [+]
      FlyBase Computed Cytological Location
      Cytogenetic map
      Evidence for location
      92F3-92F3
      Limits computationally determined from genome sequence between P{PZ}l(3)1058510585 and P{EP}SyndEP409
      Experimentally Determined Cytological Location
      Cytogenetic map
      Notes
      References
      92E12-92E14
      (determined by in situ hybridisation)
      92E14-92E15
      (determined by in situ hybridisation)
      Experimentally Determined Recombination Data
      Left of (cM)
      Right of (cM)
      Notes
      Stocks and Reagents
      Stocks (47)
      Genomic Clones (21)
      cDNA Clones (98)
       

      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 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
      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: H CG5460

      Source for database merge of
      Additional comments
      Other Comments

      Haploinsufficient locus (not associated with strong haplolethality or haplosterility).

      H protein opposes N pathway activity by acting as an adaptor between Su(H) protein and the corepressors gro and CtBP.

      H is required to repress the transcription of sim.

      H forms DNA bound complex with Su(H).

      gro and H both act to restrict neuronal cell fate in the retina.

      gro and H control different aspects of ato's complex expression pattern. H represses ato expression in territories far ahead of the morphogenetic furrow, while behind the morphogenetic furrow H ensures the timely selection of regularly spaced R8 precursors, presumably by modulating N signaling.

      The activation of Su(H) by N involves the release of Su(H) from an inhibitory complex which contains H.

      The Su(H) phenotype is epistatic to that of H during wing disc development.

      H influences the whole of the sensory organ cell lineage.

      Subcellular localisation of H protein shows a major focus of activity within the nucleus.

      Candidate gene for quantitative trait (QTL) locus determining bristle number.

      Seven EMS-induced mutations have been isolated that suppress the bristle and wing vein phenotypes of NAx-16. The mutants are apparently duplications of the H locus.

      H acts by inhibiting signal transduction rather than by promoting signal production, during both selection of microchaete precursors in the notum and vein cell differentiation in the wing. H plays no essential role during embryogenesis.

      Definitive denticle belt phenotypes (abnormal hairs and/or bristles) are foreshadowed by abnormal organisation of the actin cytoskeleton in embryonic epidermal cells.

      Structure-function analysis of H demonstrates the protein can be subdivided into the absolutely essential Su(H)-binding domain, important N- and C- terminal domains and a central antimorphic domain.

      A screen to isolate mutations that cause dominant enhancement of wing vein phenotypes associated with Egfr recovered mutations in H.

      Loss of function mutations of H and the AS-C are epistatic to Brd.

      Mutations show strong interactions with high and low selection lines, abdominal and sternopleural bristle numbers are affected. Results suggest H is a candidate for bristle number quantitative trait loci (QTL) in natural populations or is in the same genetic pathway.

      N, H and Su(H) exhibit dose-dependent genetic interactions.

      H promotes stable commitment to the sensory organ precursor cell fate by negatively regulating the activity of the N signalling pathway.

      Glutathione-S-transferase (GST) fusion proteins and the yeast two hybrid system demonstrate the H protein can inhibit the in vitro DNA binding activity of Su(H) through direct protein-protein interactions. Co-transfection assays in S2 cells demonstrate transcriptional activation driven by Su(H) is also inhibited by H.

      N receptor activity suppresses H function during adult external sense organ development. H activity insulates sensory organ precursor cells from inhibition by N. H function appears to be an integral negative regulatory element of the N signalling pathway.

      Overexpression of a wild type construct causes premature lethality and gain of function phenotypes, at ambient temperature it still rescues the dominant loss of function phenotypes. Mutant derivatives have allowed separation of a lethally mutable domain from a neural domain required for sensory organ formation.

      In vitro binding assays demonstrate the H gene product can inhibit the DNA binding of Su(H) through direct protein-protein interactions. Consistent with this, transcriptional activation driven by Su(H) in transfected S2 cells is inhibited by H. These results support a model in which H acts, in part, as a negative regulator of Su(H) activity.

      Decreasing or increasing the level of Su(H) function confers mutant phenotypes that closely resemble those associated with gain and loss of H activity, respectively.

      The phenotypes of the Abruptex class of N allele are modified by mutations at Ser, Dl, H and gro.

      Dominant lethal interaction of an allele of H with kniri-92f reported: transheterozygotes are never recovered.

      The H gene product is involved in the early nervous system differentiation and counteracts the action of other neurogenic genes.

      NM1 defines a new class of Notch allele: similarity with and lack of specificity of interaction of N- and NM1 with H, mam, gro and E(spl) suggest that the NM1 effect is due to modification in the intracellular signalling of the activated N receptor.

      Whereas H mutant alleles cause trichogen to tormogen double socket phenotype, heat shock ectopic Hairless-expression experiments produce double shaft tormogen to trichogen transformation, and bristle multiplication/loss phenotype similar to those caused by loss of function Notch mutations. Sensory organ precursor cell fate may require H to antagonise neurogenic gene functions and expression of distinct trichogen/tormogen sister cell pair may depend on asymmetry of H+ activity or their response to it. H has been cloned beginning with plasmid rescue of the insertion in HD179. Heat shock driven H+ line has been generated that rescues mutant H alleles. Mutant phenotype and expression studies suggest that neither fine spatial/temporal regulation nor high levels of H transcription are required for normal H function.

      A screen that identified Dominant suppressors of the wing phenotype associated with Dl9P and DlFE32 identified 12 new H alleles.

      In H mutants, neuronal precursors do not differentiate: H may be involved in specifying neuronal cell fate. H mutations show genetic interactions with neurogenic mutations.

      Genetic and molecular study of H function reveals that H is one of a small number of genes for which the loss of function phenotype is the failure of sensillum precursor development.

      emcD has synergistic interactions with H alleles and mutual rescuing with Dl alleles. The NAx-1, NAx-M2 and NAx-M3 alleles increase the phenotype of H alleles.

      Lack-of-function alleles of H correct ASC "Hw" phenotypes in both ectopic and normal positions.

      H is an enhancer of spl alleles of N: due to the haploinsufficiency of the H locus.

      Mutations in H cause pleiotropic phenotypes in embryonic patterns and affect several longitudinal veins.

      H dominantly suppresses the dx adult wing and ocelli phenotypes. Flies heterozygous for H and homozygous for dxENU have eyes smaller than wild type, infrequent missing or duplicated bristle phenotype and ommatidia are square wih a bristle at each corner.

      ve, vn, ci, cg, svs, ast, H, Vno and vvl belong to the vein phenotypic group (Puro, 1982, Droso. Info. Serv. 58:205--208 ) within the 'lack-of-vein' mutant class. Loss-of-function alleles at these loci remove stretches of veins in two or more longitudinal veins. Double mutations within members of this group remove all veins, have smaller, slightly lanceolate wings, no sensilla and extra chaetae. Some alleles are embryonic lethal.

      A study of the interactions between N, Dl, H and E(spl) suggest that the effects of H, Dl and E(spl) on N are allele specific and occurring at the protein level.

      Functional relationships between neurogenic genes has been studied.

      Bristles, especially postverticals and abdominals, missing in heterozygous H flies. Bristle sockets present at some sites, not at others. Expressed most distinctly on head; occipital; postvertical and ocellar bristles affected. Bristles of antennae and vibrissae show mutant phenotype much less frequently. Sockets without shafts also found on thorax, scutellum, abdominal tergites, external genitalia, wings, and legs. No shaftless sockets appear on the bracted costa of the wing. Some 40% of bristle organs located on distal part of femur differentiate neither shaft nor bract; bracts absent whenever shaft missing but present when shaft present; abnormally short shaft may be accompanied by normal-sized bract (from description of H2, by Tobler, Rothenbuhler and Nothiger (1973). Veins L4 and L5 do not reach wing margin; occasionally true of L2 also. Eyes larger than wild type; body color somewhat paler. Lees and Waddington (1942) show that trichogen cell forms a socket instead of a bristle shaft at some sites. Phenotypic expression of H responds linearly to dosage of Su(H)+ in region 35B6-10 on the left arm of chromosome 2. The number of microchaetae in H/+ flies varies from approximately 35 in the presence of a single dose of 35B6-10 to fewer than 10 in the presence of four doses (Ashburner, 1982). Interactions with other mutants studied by House (House, 1953a; House 1953b; House, 1959a; House, 1955; House, 1959b). H suppresses wing notching of N, fa, Nnd-3 and nd, enhances Ax; also H enhances eye effect of spl and removes more bristles in combination with spl (House, Von Halle). Reduction in the number of copies of the wild-type allele of H decreases the mutant phenotype of heterozygous N and Dl flies, but increase in the number of copies of the wild-type allele of H enhances the mutant phenotype of heterozygous N, Dl and E(spl) flies (Vassin, Vielmetter and Campos-Ortega, 1985). H shows some superadditive interaction with en, ci, ciW and ciD relative to degree of L4 interruption. L2 interruption augmented in combinations with ve and ri; L3 interruption augmented in combinations with ve and tt. Triploid, H/+/+, intermediate between wild type and H/+. H/H/+ most extreme type with bristles absent from head, thorax and abdomen (Gowen, 1933). Homozygous lethal. H null homozygotes die during larval and pupal stages (Bang, Hartenstein and Posakony, 1991). Animals surviving to pharate adult are completely devoid of macrochaetes and microchaetes on the head and notum, with occasional 'double sockets' remaining on the abdominal tergites. Bristles on the legs significantly resistant to loss of H+ function; many 'double sockets' and some normal bristles remain on the legs of H null homozygotes. In regions of the notum exhibiting bristle 'loss' in adult H mutants, macrochaete and microchaete primary precursor cells undetectable (Bang, Hartenstein and Posakony, 1991).

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      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.
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      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.
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      BioGRID - A database of protein and genetic interactions.
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      DRSC - Results frm RNAi screens
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      FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
      FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
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      Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
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      MIST (genetic) - An integrated Molecular Interaction Database
      MIST (protein-protein) - An integrated Molecular Interaction Database
      Synonyms and Secondary IDs (5)
      Reported As
      Symbol Synonym
      H
      (Gahr et al., 2019, Guo et al., 2019, Maier, 2019, Miller et al., 2019, Shokri et al., 2019, Smylla et al., 2019, Zacharioudaki et al., 2019, Bala Tannan et al., 2018, Gomez-Lamarca et al., 2018, Kavaler et al., 2018, Lan et al., 2018, Salazar and Yamamoto, 2018, Chan et al., 2017, Nagel et al., 2017, Praxenthaler et al., 2017, Shukla et al., 2017, Zehender et al., 2017, Zhu et al., 2017, Horvath et al., 2016, Malartre, 2016, Morimoto et al., 2016, Smylla et al., 2016, Dai et al., 2015, Praxenthaler et al., 2015, Brockmann et al., 2014, Coumailleau and Schweisguth, 2014, Fernandes et al., 2014, Gómez-Lamarca et al., 2014, Nagel and Preiss, 2014, Zacharioudaki and Bray, 2014, Dai et al., 2013, Das et al., 2013, de Celis et al., 2013.9.11, Djiane et al., 2013, Wang et al., 2013, Yu et al., 2013, Cook et al., 2012, Domanitskaya and Schüpbach, 2012, Hainaut et al., 2012, Hazelett et al., 2012, Sun et al., 2012, Troost and Klein, 2012, Guan et al., 2011, Johnson et al., 2011, Maier et al., 2011, Nagel and Preiss, 2011, Pilgram et al., 2011, Tsubota et al., 2011, Walrad et al., 2011, Zhou et al., 2011, Bardin et al., 2010, Kim et al., 2010, modENCODE Consortium et al., 2010, Saj et al., 2010, Wasbrough et al., 2010, Ayroles et al., 2009, Benítez et al., 2009, Mao and Freeman, 2009, Moshkin et al., 2009, Olswang-Kutz et al., 2009, Perkins et al., 2009.8.10, Protzer et al., 2009, Stern et al., 2009, Bejarano et al., 2008, Haussmann et al., 2008, Hayashi et al., 2008, Leevers, 2008.10.20, Maier et al., 2008, Parks et al., 2008, Protzer et al., 2008, Yatsu et al., 2008, Araujo et al., 2007, Goodfellow et al., 2007, Harding-Theobald et al., 2007.2.6, Kugler and Nagel, 2007, Martin et al., 2007, Minidorff et al., 2007, Minidorff et al., 2007, Nagel et al., 2007, Park et al., 2007, Thomas and van Meyel, 2007, Herranz et al., 2006, Koelzer and Klein, 2006, Mahoney et al., 2006, Molnar et al., 2006, Müller et al., 2006, Orme et al., 2006, Williams et al., 2006, Copley, 2005, Macdonald and Long, 2005, Nagel et al., 2005, Tang et al., 2005, Wech and Nagel, 2005, Wesley and Mok, 2003, Schreiber et al., 2002, Gim et al., 2001, Weinmaster, 1997)
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