FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\Hmr
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General Information
Symbol
Dmel\Hmr
Species
D. melanogaster
Name
Hybrid male rescue
Annotation Symbol
CG1619
Feature Type
FlyBase ID
FBgn0001206
Gene Model Status
Stock Availability
Gene Summary
Hybrid male rescue (Hmr) encodes a protein required to repress transposable element and satellite DNA expression. It also has a gain-of-function phenotype of causing lethality in F1 male hybrids between D. melanogaster and D. simulans. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
Also Known As

BcDNA:LD22117

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
1-32
RefSeq locus
NC_004354 REGION:10590920..10595855
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (13 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from physical interaction with FLYBASE:Lhr; FB:FBgn0034217
Terms Based on Predictions or Assertions (0 terms)
Biological Process (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
Cellular Component (7 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
inferred from direct assay
located_in nucleolus
inferred from direct assay
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000956409
inferred from biological aspect of ancestor with PANTHER:PTN000956409
Protein Family (UniProt)
-
Summaries
Gene Snapshot
Hybrid male rescue (Hmr) encodes a protein required to repress transposable element and satellite DNA expression. It also has a gain-of-function phenotype of causing lethality in F1 male hybrids between D. melanogaster and D. simulans. [Date last reviewed: 2019-03-07]
Gene Group (FlyBase)
MADF-BESS DOMAIN TRANSCRIPTION REGULATORS -
The MADF-BESS domain transcription regulators include a variety of activators and corepressors and chromatin modifying proteins. BESS domains are often present in combination with MADF domains. The BESS domain mediates protein-protein interactions. (Adapted from FBrf0155780 and FBrf0224045).
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
hmr: hybrid male rescue
Rescues D. melanogaster / simulans hybrid males that would not survive in the absence of the gene. Hybrid males that carry both hmr and a duplication (thought to be hmr+) are lethal, so the rescue is considered recessive. The hybrids are sterile. Hybrid females from the reciprocal cross show low viability. D. melanogaster / mauritania and D. melanogaster / sechellia hybrid males also rescued. The rescue of D. melanogaster / simulans and D. melanogaster / mauritania males by hmr almost complete at 18; at 25 D. melanogaster / mauritania male rescue is good, but D. melanogaster / simulans males is poor at this temperature. At 18 only one third of the D. melanogaster / sechellia males are rescued.
Summary (Interactive Fly)

chromatin factor - contains BESS motif found in boundary element proteins - along with Hybrid male rescue interacts with HP1 to repress transcripts from satellite DNAs and many families of transposable elements - gain-of-function phenotype causing lethality in F1 male hybrids between D. melanogaster and D. simulans

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

Please see the JBrowse view of Dmel\Hmr 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 Q86CW5)

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 5.51

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0071511
4696
1413
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
FBpp0071440
158.6
1413
10.08
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\Hmr 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).

-0.23

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

Comment: maternally deposited

Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
inferred from direct assay
located_in nucleolus
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Hmr 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
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 7 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 16 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Hmr
Transgenic constructs containing regulatory region of Hmr
Aberrations (Deficiencies and Duplications) ( 15 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (1)
1 of 14
Yes
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (0)
Mus musculus (laboratory mouse) (0)
Xenopus tropicalis (Western clawed frog) (0)
Danio rerio (Zebrafish) (6)
1 of 14
Yes
No
1 of 14
Yes
No
1 of 14
Yes
No
Caenorhabditis elegans (Nematode, roundworm) (4)
2 of 14
Yes
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (13)
1 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (1)
1 of 13
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Hmr. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (27)
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Interaction
    References
    Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    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

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    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
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    FlyBase
    External Links
    External Data
    Linkouts
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    X
    Recombination map
    1-32
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    9D3-9D3
    Limits computationally determined from genome sequence between P{EP}PPP4R2rEP307 and P{EP}rasEP1235&P{EP}rasEP1427
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    9E-9E
    (determined by in situ hybridisation)
    Location based on the correlation between cytogenetic and meiotic map positions in the middle region of the X chromosome, and its genetic map position of 1-31.84 as reported in FBrf0047061).
    Method of mapping is unspecified.
    Hmr maps distal to the distal In(1)AB breakpoint.
    Experimentally Determined Recombination Data
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (11)
    Genomic Clones (20)
    cDNA Clones (30)
     

    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
      RNAi and Array Information
      Linkouts
      DRSC - Results frm RNAi screens
      Antibody Information
      Laboratory Generated Antibodies
       
      Commercially Available Antibodies
       
      Cell Line Information
      Publicly Available Cell Lines
       
        Other Stable Cell Lines
         
        Other Comments

        The Hmr and Lhr proteins form a heterochromatic complex with Su(var)205 in D. melanogaster. Hmr and Lhr are required to repress transcripts from satellite DNAs and many families of transposable elements. They are also required to help regulate the length of telomeres (which in Drosophila are composed of domesticated transposable elements).

        The Dsim\Lhr ortholog of Lhr does not appear to affect satellite DNA transcription in D. simulans, but it does have a role in repressing transcription of transposable elements in this species.

        Upregulation of transposable element transcription is seen in hybrids between D. melanogaster and D. simulans, but this is unlikely to be the direct cause of hybrid lethality.

        The Hmr and Lhr proteins form a centromeric complex in D. melanogaster which is required for proper chromosome segregation during mitosis. Both an increase and a decrease in complex levels result in mitotic defects, indicating that this function is extremely dose sensitive. Alteration of the levels of the complex also result in an increase in transcription from transposable elements.

        The orthologous D. simulans Dsim\Hmr and Dsim\Lhr proteins also bind each other in coimmunoprecipitation experiments and also localise to the centromere (although Dsim\Hmr also shows a prominent non-centromeric localisation).

        The level of Hmr expression in D melanogaster is substantially higher than the expression of the orthologous Dsim\Hmr gene in D. simulans. Conversely, the expression of the D. simulans Dsim\Lhr gene is higher than the expression of the orthologous Lhr gene in D. melanogaster. Hybrids derived from a cross between D. melanogaster females and D. simulans males thus have an elevated amount of the Hmr-Lhr protein complex compared to the parent species, and the complex is delocalised, being bound to numerous interbands along all chromosome arms in the hybrids. Hybrid males and females show a massive increase in transcription from transposable elements, but as the effect is not sex-specific, this is presumably not the the main cause of the lethality of hybrid males.

        Hmr is required for wild-type levels of viability and fertility of D.melanogaster females.

        Dsim\Lhr has functionally diverged in D.simulans and interacts with Hmr, which has functionally diverged in D.melanogaster, to cause lethality in F1 hybrid males.

        Mutations in the D.melanogaster gene Hmr, along with unidentified polymorphic factors, rescue the agametic phenotype of F1 hybrid females derived from crosses of D.melanogaster females to either D.simulans or D.mauritiana males. F1 hybrid males from these crosses are fully sterile. The F1 hybrid females produce small numbers of progeny in backcrosses to D.melanogaster males, the low fecundity being caused by incomplete rescue of oogenesis as well as zygotic lethality.

        Hmr has a role in inter-species hybrid incompatability.

        The wild-type product of Hmr is neither necessary nor sufficient for embryonic inviability in hybrids between D.melanogaster and its sibling species. Hmr does, however, appear to lower the viability of hybrid larvae. This data suggests that Hmr acts as a gain-of-function poison in hybrids.

        Mapping and characterisation of Hmr.

        Hybrid female progeny resulting from crossing D.simulans females to D.melanogaster males die as embryos. Hybrid males from the reciprocal cross die as late larvae, unless the male parent is mutant at Dsim\Lhr, or the female at Hmr. Hybrid males from D.simulans attached X females and D.melanogaster males are lethal at both embryonic and larval stages, but are rescued to viability by the Hmr mutant in the male parent and the Dsim\mhr mutant in the female parent.

        Relationship to Other Genes
        Source for database merge of

        Source for merge of: Hmr BcDNA:LD22117

        Additional comments

        A Dsim\Hmr+ transgene has no phenotypic effect in D.melanogaster/D.simulans and D.melanogaster/D.mauritiana hybrids and a Dmau\Hmr+ transgene has no phenotypic effect in D.melanogaster/D.simulans and D.melanogaster/D.mauritiana hybrids, strongly supporting the conclusion that the Hmr gene has functionally diverged between the D.melanogaster and sibling-species (D.simulans and D.mauritiana) lineages, to cause F1 hybrid incompatibility between these species. Phylogenetic analysis shows that Hmr has diverged extensively in the D.melanogaster lineage, but extensive divergence has also been found in the sibling-species lineage. Together, the findings implicate over 13% of the amino acids encoded by Hmr as candidates for causing hybrid incompatibility. The exceptional level of divergence at Hmr cannot be explained by neutral processes, as phylogenetic methods and population genetic analyses show that the elevated amino-acid divergence in both lineages is due to positive selection in the distant past - at least one million generations ago.

        It is possible that "anon-AE003451.1" should be merged with " BcDNA:LD22117 " to make a single gene.

        sesB and Ant2 do not correspond to Hmr; Hmr maps distal to both genes and neither sesB mutant alleles nor transformants carrying extra copies of sesB and Ant2 have any effect on interspecific viability.

        Source for merge of Hmr BcDNA:LD22117 was sequence comparison ( date:030514 ).

        Nomenclature History
        Source for database identify of
        Nomenclature comments
        Etymology
        Synonyms and Secondary IDs (9)
        Reported As
        Secondary FlyBase IDs
        • FBgn0027511
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 72 )
        Sequence Crossreferences
        NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
        GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
        RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
        UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
        UniProt/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
        InterPro - A database of protein families, domains and functional sites
        KEGG Genes - Molecular building blocks of life in the genomic space.
        MARRVEL_MODEL - MARRVEL (model organism gene)
        Linkouts
        BioGRID - A database of protein and genetic interactions.
        Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
        DroID - A comprehensive database of gene and protein interactions.
        DRSC - Results frm RNAi screens
        Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
        FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
        FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
        Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
        Flygut - An atlas of the Drosophila adult midgut
        Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
        MIST (protein-protein) - An integrated Molecular Interaction Database
        References (116)