Open Close
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 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]
Also Known As

BcDNA:LD22117

Key Links
Genomic Location
Cytogenetic map
Sequence location
X:10,590,920..10,595,855 [-]
Recombination map

1-32

RefSeq locus
NC_004354 REGION:10590920..10595855
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (13 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000496516
(assigned by GO_Central )
Biological Process (5 terms)
Terms Based on Experimental Evidence (4 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
inferred from biological aspect of ancestor with PANTHER:PTN000496516
(assigned by GO_Central )
Cellular Component (6 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000496516
(assigned by GO_Central )
Protein Family (UniProt)
-
Summaries
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.
Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\Hmr or 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
Comments on Gene Model

Gene model reviewed during 5.51

Sequence Ontology: Class of Gene
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
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
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

Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
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\Hmr 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
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 7 )
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 Hmr
Transgenic constructs containing regulatory region of Hmr
Deletions and Duplications ( 15 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Sterility
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (3)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
No
Rattus norvegicus (Norway rat) (1)
1 of 13
Yes
No
Xenopus tropicalis (Western clawed frog) (1)
1 of 12
Yes
No
Danio rerio (Zebrafish) (1)
1 of 15
Yes
No
Caenorhabditis elegans (Nematode, roundworm) (1)
1 of 15
Yes
No
Arabidopsis thaliana (thale-cress) (5)
1 of 9
Yes
No
1 of 9
Yes
No
1 of 9
Yes
No
1 of 9
Yes
No
1 of 9
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (1)
2 of 15
Yes
No
Schizosaccharomyces pombe (Fission yeast) (1)
1 of 12
Yes
No
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091901AO )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091500RJ )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( None identified )
No non-Dipteran orthologies identified
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X00K9 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
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) (2)
2 of 10
2 of 10
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
    Summary of Genetic Interactions
    esyN Network Diagram
    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.
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    External Data
    Linkouts
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    X
    Recombination map

    1-32

    Cytogenetic map
    Sequence location
    X:10,590,920..10,595,855 [-]
    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 (12)
    Genomic Clones (20)
    cDNA Clones (33)
     

    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 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 ).

      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.

      Origin and Etymology
      Discoverer
      Etymology
      Identification
      External Crossreferences and Linkouts ( 69 )
      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/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
      InterPro - A database of protein families, domains and functional sites
      KEGG Genes - Molecular building blocks of life in the genomic space.
      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
      FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
      FlyMine - An integrated database for Drosophila genomics
      InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
      MIST (protein-protein) - An integrated Molecular Interaction Database
      Synonyms and Secondary IDs (9)
      Datasets (0)
      Study focus (0)
      Experimental Role
      Project
      Project Type
      Title
      References (101)