General Information
Symbol
Dmel\dsx
Species
D. melanogaster
Name
doublesex
Annotation Symbol
CG11094
Feature Type
FlyBase ID
FBgn0000504
Gene Model Status
Stock Availability
Gene Snapshot
In progress.Contributions welcome.
Also Known As
doublesex, Dmdsx
Genomic Location
Cytogenetic map
Sequence location
3R:7,924,323..7,967,408 [-]
Recombination map
3-48
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
GO Summary Ribbons
Families, Domains and Molecular Function
Gene Group Membership (FlyBase)
Protein Family (UniProt, Sequence Similarities)
-
Summaries
Gene Group Membership
DMRT TRANSCRIPTION FACTORS -
The DMRT (doublesex- and mab-3-related transcription factor) proteins are dimeric sequence-specific DNA binding proteins that regulate transcription. They are characterized by a DM domain containing an intertwined C2HC and HCC2 zinc binding sites. DMRT proteins have been associated with the regulation of sex-determination genes. (Adapted from PMID:17605809).
UniProt Contributed Function Data
Controls somatic sexual differentiation. Binds directly and specifically to the FBE (fat body enhancer) of the yolk protein 1 and 2 genes (Yp1 and Yp2). This enhancer is sufficient to direct the female-specific transcription characteristic of the Yp genes in adult fat bodies. Involved in regulation of male-specific expression of takeout in brain-associated fat body.
(UniProt, P23023)
Phenotypic Description from the Red Book (Lindsley and Zimm 1992)
dsx: double sex (B. J. Taylor)
The dsx gene regulates sexual differentiation of somatic tissues. Null alleles convert chromosomally male and female flies into sterile intersexes of similar phenotype. Dominant alleles (e.g., dsxD, dsxM, dsxT) transform females into intersexes when heterozygous with a normal allele, and into phenotypic males when homozygous or heterozygous with a dsx-null allele or deficiency, but they have no effect in males. Most alleles at dsx affect both sexes; however, some alleles affect only one sex. The recessive allele dsx11 converts males into intersexes and is complemented by dominant dsx alleles and recessive alleles that affect only females (dsx22) (Baker and Ridge; Nothiger et al., 1987). Double-mutant combinations of dsx null mutations with loss-of-function alleles at tra, tra2, and ix result in a doublesex phenotype (Mukherjee and Hildreth, 1971, Genetica 42: 338-52; Baker and Ridge; Nothiger et al., 1987). Double-mutant combinations of dsxD/+ with null alleles of tra and tra2 convert females into phenotypic males or with ix into more male-like intersexes (Baker and Ridge; Nothiger et al., 1987). The dose of dsx alleles can alter the phenotype; triploid female flies dsxD/+/+ are sterile and with a weak external dsx phenotype (Gowen and Fung, 1957; Nothiger et al., 1987); diploid female flies that are dsxD/+, but also carry a dsx+ duplication Tp(3;Y)P92 are sterile but female in appearance (Nothiger et al., 1987). Germline sexual differentiation is not dependent on dsx+ function; only the chromosomal constitution determines the sex of transplanted dsxM/+, dsx1, dsxD/+, and dsxD/dsx1 germ cells (Nothiger, Roost, and Schupbach, 1980, DIS 55: 118; Schupbach, 1982, Dev. Biol. 89: 117-27). The dsx+ gene does not appear to encode any vital functions (Baker and Ridge). The normal body size differences between male and female flies is maintained in dsx-null mutants (Hildreth) and in females heterozygous for dsxD/+, dsxD/dsx1 (Fung and Gowen, 1957; Baker and Ridge) and dsxM/+ (Nothiger et al., 1987). The sexcomb bristles on the prothoracic basitarsus in both sexes of dsx-null homozygotes (Hildreth; Mukherjee, and Hildreth; Baker and Ridge) and female dsxD/dsx1 (Nothiger et al., 1987) are intermediate in number, morphology, and position compared with the sexcomb bristles in normal males and the transverse row bristles in normal females. The central sexcomb bristle is retained in dsx-null mutants (Hildreth). In dsx-null mutants, the pigmentation of the fifth tergite is intermediate between the completely pigmented male and the posteriorly pigmented female tergite, whereas the sixth tergite is darkly pigmented (Hildreth; Baker, and Ridge). Female flies that are dsxD/+ or dsxM/+ are similar to dsx homozygotes (Fung and Gowen, 1957; Duncan and Kaufman, 1975, Genetics 80: 733-52; Baker and Ridge; Nothiger et al., 1987). Male dsx flies have a seventh tergite and sternite with bristles (Hildreth; Baker, and Ridge). Female flies heterozygous for dominant alleles and either dsx1 or dsx deficiencies have the male number of tergites and sternites with the male pattern of pigmentation (Duncan and Kaufman, 1975; Baker and Ridge; Nothiger et al., 1980; Nothiger et al., 1987). By clonal analysis, the action of dsx has been shown to be cell autonomous in the differentiation of the sexcombs and pigmentation of the abdominal tergites; dsx+ is required until the end of the larval period for the proper sexual differentiation of the sexcombs and into the pupal period, close to the time of the termination of divisions of the abdominal histoblasts, for proper sexual differentiation of the abdominal histoblasts and for proper sexual differentiation of the abdomen (Baker and Ridge). Both male and female genitalia are formed in dsx null mutant flies and in female flies heterozygous for dominant alleles (Fung and Gowen, 1957; Hildreth, 1965; Epper, 1981, Dev. Biol. 88: 104-14; Nothiger et al., 1987); a second penis differentiates with a reduced aedeagus and parameres within the female vaginal area (Hildreth). In dsxD/+ females, the development of the female genitalia and second penis are very similar to that of dsx-null flies, whereas in dsxM/+ females the female genitalia are more severely reduced (Gowen and Fung, 1975; Baker and Ridge; Nothiger et al., 1980; Epper, 1981; Nothiger et al., 1987). Male genitalia from dsx null flies and females heterozygous for dsx dominant alleles contain all elements except a basal apodeme but other external structures such as the penis and accessory elements are reduced and not as well formed (Fung and Gowen, 1957; Hildreth; Epper, 1981). The internal duct systems develop but can vary between dual female and male ducts and a single poorly differentiated duct (Hildreth); a similar range of phenotypes for the internal ducts is found in dsxD/+ (Fung and Gowen, 1957) and dsxM/+ (Nothiger et al., 1980). Based on fate mapping and analysis of the morphogenesis of the dsxD/+ genital disc, the female genitalia and second penis are generated from the female genital primordium, and the male genitalia from the male genital primordium and the production of both types of genitalia in dsx flies results from derepression of both genital primordia (Epper, 1981; Epper, 1983, Wilhelm Roux's Arch. Dev. Biol. 192: 280-84). The intersexual analia differentiate as lateral plates, which are smaller than normal male lateral anal plates and do not have the ventral anal plate found in females (Hildreth; Baker, and Ridge; Epper, 1981). The bristle pattern is rather male-like but with one clearly identifiable long dorsal bristle that is female (Epper, 1981). The gonads are often rudimentary, but occasionally female dsx-null as well as dsxD/+ flies have well developed ovaries and eggs (Hildreth, Fung, and Gowen, 1957; Schupbach, 1982; Bownes, Dempster, and Blair, 1983, J. Embryol. Exp. Morph. 75: 241-57) whereas in male dsx-null flies, the gonads are poorly developed and no sperm are formed (Hildreth; Schupbach, 1982). Female and male dsx flies and female dsxD/+ flies make yolk protein but in amounts less than for normal females (Postelthwait, Bownes, and Jowett, 1980, Dev. Biol. 79: 379-87; Ota, Fukunaga, Kawabe, and Oishi, 1981, Genetics 99: 429-41). In dsxD/+ females less yolk protein synthesis occurs compared to normal females; little or no yolk protein mRNA is made in the rudimentary gonads, but measurements from adult flies show that from an initially low level yolk protein transcripts increase to nearly wildtype levels but without efficient conversion into protein (Bownes, Dempster, and Blair, 1983). The male-specific transcripts 316, 355a, and 355b, made by the male accessory gland, are also produced in male flies rendered intersexual by the mutation dsx11, which does not affect female flies; in addition, females that are dsxD/+ or dsxM/+ express the male-specific transcripts, although at reduced levels (Chapman and Wolfner, 1988, Dev. Biol. 126: 195-202). Females homozygous for dsx do not exhibit male courtship behaviors (McRobert and Tompkins, 1985, Genetics 111: 89-96; B. Taylor, unpublished). These dsx females do not make 7, 11 dienes, and 7 monoenes compared to normal females (Jallon, 1984, Behav. Genet. 14: 441-78); they elicit less courtship than dsx+ females from control males (McRobert and Tompkins, 1985). Female flies that are dsxD/+ or dsxM/+ also fail to express male courtship behaviors (Duncan and Kaufman, 1975; B. Taylor, unpublished). Males homozygous for dsx court but show reduced levels of courtship directed toward females and young males and greater-than-normal levels of courtship directed at mature males (McRobert and Tompkins, 1985). Mature dsx males make some pheromonal substances (Jallon, 1984) and elicit more courtship than dsx+ males from control males (McRobert and Tompkins, 1985). Female flies homozygous for dsx-null alleles or mutant for dsx dominant alleles do not make the male-specific abdominal muscle unlike their male siblings (B. Taylor, unpublished).
Gene Model and Products
Number of Transcripts
6
Number of Unique Polypeptides
3

Please see the GBrowse view of Dmel\dsx or the JBrowse view of Dmel\dsx 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_with_stop_codon_read_through ; SO:0000697
Stop-codon suppression (UAG) postulated; FBrf0216884.
Gene model reviewed during 5.44
Gene model reviewed during 5.49
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0081759
4010
549
FBtr0081760
3627
427
FBtr0081761
3238
427
FBtr0330073
3621
549
FBtr0330074
4010
572
FBtr0339710
6941
427
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
FBpp0081256
57.4
549
8.83
FBpp0081257
44.8
427
7.97
FBpp0081258
44.8
427
7.97
FBpp0303106
57.4
549
8.83
FBpp0303107
59.6
572
8.83
FBpp0308767
44.8
427
7.97
Polypeptides with Identical Sequences

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

549 aa isoforms: dsx-PA, dsx-PD
427 aa isoforms: dsx-PB, dsx-PC, dsx-PF
Additional Polypeptide Data and Comments
Reported size (kDa)
549, 427 (aa); 57.4, 44.8 (kD predicted)
Comments
male-specific carboxy-terminal end
amino-terminal end
female-specific carboxy-terminal end
female-specific
male-specific
External Data
Crossreferences
PDB - An information portal to biological macromolecular structures
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\dsx 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 (28 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Biological Process (21 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (16 terms)
CV Term
Evidence
References
Cellular Component (1 term)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR026607
(assigned by InterPro )
inferred by curator from GO:0001228
Expression Data
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: maternally deposited

RT-PCR
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: male-specific transcript only

Additional Descriptive Data
The male-specific form of dsx transcript is expressed solely in male embryos. It is expressed in male-specific somatic gonadal precursor cells.
dsx transcript is expressed in the larval and adult CNS. RT-PCR analysis using transcript-isoform-specific primers shows male- and female-specific expression.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
central nervous system | restricted

Comment: reference states 1-2 days APF

protocerebrum | restricted

Comment: reference states 1-2 days APF

subesophageal ganglion | restricted

Comment: reference states 1-2 days APF

thoracico-abdominal ganglion | restricted

Comment: reference states 1-2 days APF

neuron | subset of adult brain

Comment: Antibody detects male-specific protein.

adult fruitless pMP-e (male) lineage clone

Comment: referred to as P1. Antibody detects male-specific protein.

Additional Descriptive Data
Antibodies to the male-specific form of dsx detect protein in all male-specific somatic gonadal precursor cells in late embryo but not in the germline cells. These somatic cells are intermingled with germline cells. Male-specific dsx is expressed in all posterior somatic gonadal cells expressing eya and either Sox100B or tj. There is another population of cells that wraps around the embryonic testis at embryonic stage 17 that express Sox100B but not dsx. dsx is also detected in hub cells in stage 17 embryos. The male-specific dsx isoform is detected in cyst cells in larval and adult testis.
In the late third instar larval central nervous system, dsx protein is distributed in a relatively small number of cells in the brain lobes and ventral nerve cord. The most broad and intense dsx immunoreactivity in the CNS is observed in pupae 1-2 days APF. Labeled cells in each brain hemisphere include 2 anterior-dorsal neurons in the superior protocerebrum, 2-3 lateral subesophageal neurons, 1 neuron located medially in the ventral-most part of the subesophageal ganglion, and two groups of 30-50 cells each located posteriorly and dorsally to the mushroom body calyx; about 20-30 cells of these last two groups are non-neuronal. In the ventral nerve cord, labeled cells include 18-24 neurons per side in the prothoracic and metathoracic ganglia; another 38-42 neurons in the thoracic ganglia; and 200-300 neurons in the abdominal ganglia. A similar but fainter pattern is observed in later pharate adults (3-4 days APF), and most dsx-expressing cells observed in pupal CNS are also observed in the adult CNS, with much fainter immunoreactivity in female adults than in male adults.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
Reporter: P{GMR40F04-GAL4}
Stage
Tissue/Position (including subcellular localization)
Reference
Stage
Tissue/Position (including subcellular localization)
Reference
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: TI{GAL4}dsxGAL4
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: TI{GAL4}dsxKI.GAL4
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\dsx 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) 11-12
  • Stages(s) 13-16
Alleles, Insertions, Transgenic Constructs and Phenotypes
Classical and Insertion Alleles ( 86 )
For All Classical and Insertion Alleles Show
 
Allele of dsx
Class
Mutagen
Associated Insertion
Stocks
Known lesion
    Yes
      0
      --
        0
        Yes
          0
          --
            0
            --
              0
              --
                0
                --
                  0
                  --
                    0
                    --
                    Other relevant insertions
                    Transgenic Constructs ( 26 )
                    For All Alleles Carried on Transgenic Constructs Show
                    Transgenic constructs containing/affecting coding region of dsx
                    Allele of dsx
                    Mutagen
                    Associated Transgenic Construct
                    Stocks
                    Transgenic constructs containing regulatory region of dsx
                    GAL4 construct
                    Name
                    Expression Data
                    Deletions and Duplications ( 35 )
                    Partially disrupted in
                    Summary of 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 cuticle & abdominal segment 5 | female
                    adult cuticle & abdominal segment 6 | female
                    Orthologs
                    Human Orthologs (via DIOPT v7.1)
                    Homo sapiens (Human) (8)
                    Species\Gene Symbol
                    Score
                    Best Score
                    Best Reverse Score
                    Alignment
                    Complementation?
                    Transgene?
                    3 of 15
                    Yes
                    Yes
                    3 of 15
                    Yes
                    Yes
                    2 of 15
                    No
                    No
                    2 of 15
                    No
                    No
                    2 of 15
                    No
                    No
                    2 of 15
                    No
                    No
                    2 of 15
                    No
                    Yes
                    1 of 15
                    No
                    Yes
                    Model Organism Orthologs (via DIOPT v7.1)
                    Mus musculus (laboratory mouse) (10)
                    Species\Gene Symbol
                    Score
                    Best Score
                    Best Reverse Score
                    Alignment
                    Complementation?
                    Transgene?
                    3 of 15
                    Yes
                    Yes
                    3 of 15
                    Yes
                    Yes
                    2 of 15
                    No
                    No
                    2 of 15
                    No
                    No
                    2 of 15
                    No
                    No
                    2 of 15
                    No
                    No
                    2 of 15
                    No
                    Yes
                    1 of 15
                    No
                    Yes
                    1 of 15
                    No
                    Yes
                    1 of 15
                    No
                    Yes
                    Rattus norvegicus (Norway rat) (10)
                    3 of 13
                    Yes
                    Yes
                    2 of 13
                    No
                    No
                    2 of 13
                    No
                    No
                    2 of 13
                    No
                    No
                    2 of 13
                    No
                    No
                    2 of 13
                    No
                    Yes
                    1 of 13
                    No
                    No
                    1 of 13
                    No
                    Yes
                    1 of 13
                    No
                    Yes
                    1 of 13
                    No
                    Yes
                    Xenopus tropicalis (Western clawed frog) (4)
                    1 of 12
                    Yes
                    No
                    1 of 12
                    Yes
                    No
                    1 of 12
                    Yes
                    No
                    1 of 12
                    Yes
                    No
                    Danio rerio (Zebrafish) (5)
                    2 of 15
                    Yes
                    No
                    2 of 15
                    Yes
                    No
                    2 of 15
                    Yes
                    No
                    2 of 15
                    Yes
                    No
                    1 of 15
                    No
                    Yes
                    Caenorhabditis elegans (Nematode, roundworm) (11)
                    3 of 15
                    Yes
                    No
                    3 of 15
                    Yes
                    Yes
                    3 of 15
                    Yes
                    Yes
                    2 of 15
                    No
                    No
                    2 of 15
                    No
                    No
                    2 of 15
                    No
                    Yes
                    2 of 15
                    No
                    Yes
                    2 of 15
                    No
                    Yes
                    1 of 15
                    No
                    Yes
                    1 of 15
                    No
                    Yes
                    1 of 15
                    No
                    Yes
                    Arabidopsis thaliana (thale-cress) (0)
                    No orthologs reported.
                    Saccharomyces cerevisiae (Brewer's yeast) (0)
                    No orthologs reported.
                    Schizosaccharomyces pombe (Fission yeast) (0)
                    No orthologs reported.
                    Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG09190HIG )
                    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 yakuba
                    Drosophila yakuba
                    Drosophila ananassae
                    Drosophila pseudoobscura pseudoobscura
                    Drosophila persimilis
                    Drosophila willistoni
                    Drosophila willistoni
                    Drosophila virilis
                    Drosophila mojavensis
                    Drosophila grimshawi
                    Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG09150BSI )
                    Organism
                    Common Name
                    Gene
                    Multiple Dmel Genes in this Orthologous Group
                    Musca domestica
                    House fly
                    Lucilia cuprina
                    Australian sheep blowfly
                    Mayetiola destructor
                    Hessian fly
                    Aedes aegypti
                    Yellow fever mosquito
                    Culex quinquefasciatus
                    Southern house mosquito
                    Culex quinquefasciatus
                    Southern house mosquito
                    Culex quinquefasciatus
                    Southern house mosquito
                    Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0QEH )
                    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
                    Tribolium castaneum
                    Red flour beetle
                    Pediculus humanus
                    Human body louse
                    Rhodnius prolixus
                    Kissing bug
                    Acyrthosiphon pisum
                    Pea aphid
                    Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0E24 )
                    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
                    Ixodes scapularis
                    Black-legged tick
                    Ixodes scapularis
                    Black-legged tick
                    Ixodes scapularis
                    Black-legged tick
                    Ixodes scapularis
                    Black-legged tick
                    Ixodes scapularis
                    Black-legged tick
                    Stegodyphus mimosarum
                    African social velvet spider
                    Tetranychus urticae
                    Two-spotted spider mite
                    Tetranychus urticae
                    Two-spotted spider mite
                    Tetranychus urticae
                    Two-spotted spider mite
                    Tetranychus urticae
                    Two-spotted spider mite
                    Tetranychus urticae
                    Two-spotted spider mite
                    Tetranychus urticae
                    Two-spotted spider mite
                    Tetranychus urticae
                    Two-spotted spider mite
                    Tetranychus urticae
                    Two-spotted spider mite
                    Tetranychus urticae
                    Two-spotted spider mite
                    Daphnia pulex
                    Water flea
                    Daphnia pulex
                    Water flea
                    Daphnia pulex
                    Water flea
                    Daphnia pulex
                    Water flea
                    Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
                    No non-Arthropod Metazoa orthologies identified
                    Human Disease Model Data
                    FlyBase Human Disease Model Reports
                      Alleles Reported to Model Human Disease (Disease Ontology)
                      Download
                      Models ( 0 )
                      Allele
                      Disease
                      Evidence
                      References
                      Interactions ( 0 )
                      Allele
                      Disease
                      Interaction
                      References
                      Comments ( 0 )
                       
                      Human Orthologs (via DIOPT v7.1)
                      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
                      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 look at the Interaction Group reports for full details of the physical interactions
                      protein-protein
                      Interacting group
                      Assay
                      References
                      RNA-protein
                      Interacting group
                      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
                      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.
                      Pathways
                      Gene Group - Pathway Membership (FlyBase)
                      External Data
                      Linkouts
                      Genomic Location and Detailed Mapping Data
                      Chromosome (arm)
                      3R
                      Recombination map
                      3-48
                      Cytogenetic map
                      Sequence location
                      3R:7,924,323..7,967,408 [-]
                      FlyBase Computed Cytological Location
                      Cytogenetic map
                      Evidence for location
                      84E5-84E6
                      Limits computationally determined from genome sequence between P{EP}EP3060EP3060 and P{PZ}grn05930
                      Experimentally Determined Cytological Location
                      Cytogenetic map
                      Notes
                      References
                      84E1-84E2
                      (determined by in situ hybridisation)
                      Experimentally Determined Recombination Data
                      Location
                      3-48.1
                      Left of (cM)
                      Right of (cM)
                      Notes
                      Stocks and Reagents
                      Stocks (38)
                      Genomic Clones (32)
                      cDNA Clones (24)
                       

                      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)
                      BDGP DGC clones
                      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
                       
                      Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
                      Other Information
                      Relationship to Other Genes
                      Source for database identify of
                      Source for identity of: dsx CG11094
                      Source for database merge of
                      Additional comments
                      The ability of the dsx proteins to restore V-ray formation to a Cele\mab-3 mutant is studied. The male specific splice form of dsx can restore V rays, essentially as well as wild type Cele\mab-3 can. The female specific splice form has no effect.
                      Other Comments
                      dsx is necessary for the differentiation of both male and female specific adult cuticular structures.
                      "DsxF" protein prevents the induction of dpp by hh in the repressed male primordium of female genital discs, whereas "DsxM" protein blocks the wg pathway in the repressed female primordium of male genital discs. "DsxF" protein is continuously required during female development to prevent activation of dpp in the repressed male primordium and during pupation for female genital cytodifferentiation. In males, "DsxM" is not continuously required during larval development to block the wg signaling pathway in the female genital primordium, and it does not appear to be needed during pupation for male genital cytodifferentiation.
                      Ectopic somatic expression of the female product of tra is sufficient to feminise XY germ cells. This feminisation depends upon the tra2 gene, but does not seem to require a functional dsx gene. However, feminisation of XY germ cells by the female product of tra can be blocked by the male form of dsx protein.
                      dsx is capable of repressing Dgri\Yp1 and Dgri\Yp2 in D.melanogaster males.
                      The female dsx protein plays an important role in sexual behaviour.
                      her acts together with the last genes in the determination hierarchy, dsx and ix, to control female sexual differentiation.
                      The RS domain of U2af38 is not required for enhancer-dependent splicing of dsx in vitro.
                      Sex determining genes tra, ix and dsx have no role in regulating the template organisation of the X chromosome(s) for dosage compensation.
                      DNA binding properties of purified protein dimers to dsxA, a specific DNA regulation site, is investigated; protein binding to dsxA is indistinguishable.
                      The dsx splicing enhancer contains A/C-rich splicing enhancer (ACE) motifs. A single copy of the repeat element strongly enhances splicing of vertebrate splice sites in vertebrate cells.
                      Two oligomerisation domains in male- and female-specific dsx are identified by yeast two-hybrid interaction assays and in vitro physical studies. Each protein has two oligomerisation domains; one sex independent, the other sex specific. The common function of the two domains is to oligomerize the full length protein and their specialised function is to form a dimeric DNA binding unit and a sex-specific transcriptional activation or repression unit.
                      The physical characteristics of the dsx proteins have been studied using mobility shift assays.
                      The preferred target site for dsx binding has been determined using affinity selection of random oligonucleotides and found to be a sequence with dyad symmetry, suggesting that dsx binds to its target sequence in a dimeric form. Two independent dimerization domains in the amino terminal and carboxy terminal regions of female and male specific dsx proteins have been identified and mapped using the yeast two-hybrid technique. dsx proteins expressed in the fly exist in a multimeric form.
                      fl(2)d function is necessary for the female-specific splicing of tra pre-mRNA, but not for the female-specific splicing of dsx pre-mRNA.
                      Sequences of the dsx and Dvir\dsx splicing enhancers are highly divergent except for the presence of nearly identical 13 nucleotide repeat elements (that are predominantly single stranded) and a stretch of nucleotides at the 5' and 3' ends of the enhancers. Organisation of sequences within the splicing enhancers results in a structure in which each of the repeat elements is single stranded and therefore accessible for specific recognition by the RNA binding domain of tra2.
                      Both HeLa and Kc cell nuclear extracts have been used for UV cross-linking experiments to determine which proteins bind to dsxRE as part of the native tra- and tra2-dependent dsx enhancer complex (dsxEC). Rbp1 and SRp30 have been identified that bind the 13-nucleotide repeats and purine rich element (PRE), respectively, of the dsx repeat element (dsxRE).
                      dsx mutant males are reproductively abnormal. The abnormality may stem from sexual differentiation defects in CNS, PNS or both.
                      Site directed mutagenesis, protein binding and germline transformation experiments identify and characterise the activity of a simple mini-enhancer from the fat body enhancer (FBE region) consisting of a single binding site (dsxA) for the dsx protein and two others for other regulatory proteins (slbo and ref1). One copy of this enhancer is sufficient to direct the sex and fat body specificities of Yp1 transcription.
                      Fragments of normally cis-spliced ftz pre-mRNA substrates are trans spliced in mammalian nuclear extracts. Trans splicing is promoted by a constitutively active splicing enhancer located downstream of a 3' splice site. SR proteins also promote the functional interaction of 5' and 3' splice sites in trans.
                      The ix product is required to function with the female-specific product of dsx to implement appropriate female sexual differentiation into diplo-X individuals.
                      The RNA target sequences recognised by Rbp1 have been determined using the in vitro selection approach and were found within the repeat region and in the purine rich region polypyrimidine tract of the regulated female specific 3' splice site of dsx. The Rbp1 protein can activate female specific splicing of dsx in vivo by recognising target sequences present within the pre-mRNA.
                      Female-specific expression of genes in the germline is dependent on a somatic signalling pathway which requires the sex-non-specific tra2 but not the sex-specific tra and dsx.
                      Regulated alternative splicing of dsx pre-mRNA requires the dsxRE splicing enhancer, dsx repeat element. The activity of dsxRE requires tra and tra2 and one or more general splicing factors. A purine rich enhancer (PRE) sequence within the RE has been identified, this element functionally synergises with the dsxRE and is required for specific binding of tra2 to the dsxRE. Results demonstrate that positive control of dsx pre-mRNA splicing requires tra- and tra2- dependent assembly of a multiprotein complex on at least two distinct enhancer elements. The dsx repeats R1-5 and the PRE are distinct constitutive splicing enhancer elements.
                      dsx function is required to direct the development of the genital muscles acting in wild type to repress the development of muscles of the inappropriate sex.
                      her- has no effect on dsx splicing.
                      Transcript levels from the dsx gene are not affected by nutrition.
                      The choice of female identity in the germ line is dependent upon a somatic signalling pathway that requires the sex-non-specific tra2 gene but not the sex specific genes tra and dsx.
                      The sex-specific requirement of sov in gonadal development is controlled by the somatic sex regulatory genes tra, tra2 and dsx.
                      tra and dsx control early inductive signals that determine the sex of XX germ cells.
                      dsx does not appear to materially regulate male sexual behaviour.
                      In vitro splicing reactions demonstrate the dsx repeat element (dsxRE) can act as a constitutive splicing enhancer indistinguishable from the purine-rich elements. dsxRE exhibits both constitutive and regulated activities depending on its distance from the 3' splice site.
                      In vitro mutagenesis of dsx binding sites demonstrates that in males the dsx gene product acts to directly repress transcription of the yolk genes and in females the dsx gene product activates transcription by acting at the same sites in the fat body enhancer (FBE) driving expression of Ecol\lacZ. Through the male and female dsx proteins the sexual differentiation pathway is connected to a target gene by acting directly, but with opposite effects, on the gene.
                      Both the male-specific and female-specific dsx proteins share and depend upon the same DNA binding domain for function in vivo, suggesting that both proteins bind to, but differentially regulate, a common set of genes in both sexes.
                      Results of ectopic expression of the male version of the dsx product provide evidence for a role for the male dsx protein in activation of male differentiation as well as repression of female differentiation.
                      The genetic hierarchy regulating female germ-line sex determination includes tra, tra2, dsx, fu, otu, ovo, snf and Sxl.
                      Female specific splicing of dsx is regulated by tra and tra2, which recruit general, serine/arginine-rich splicing factors to a regulatory element located downstream of a female-specific 3' splice site.
                      The M2 exon sequence of mouse IgM can stimulate the splicing of the dsx female specific intron, splicing of this intron does not usually occur due to a suboptimal pyrimidine stretch within the 3' splice site.
                      Transfection analysis in Kc cells with dsx minigene constructs identified 6 copies of a 13 nucleotide sequence in the female-specific fourth exon, that act as cis elements for female-specific splicing of dsx pre-mRNA. UV crosslinking identified tra and tra2 gene products binding to these 13 nucleotide seuqences.
                      The choice of the sexual pathway taken by sex specific neuroblasts depends on the expression of dsx.
                      dsx is a known sex determining gene, dsx does not direct the development of sexually dimorphic skeletal muscles.
                      An in vitro splicing system to study the mechanism involved in positive control of dsx female specific splicing by tra and tra2 is used in HeLa cell nuclear extracts.
                      The male and female products of dsx when expressed in E.coli bind specifically to the fat body enhancer (FBE) of Yp1 and Yp2. This demonstrates a direct interaction between the sex determination hierarchy and a target gene.
                      tra2 produced in E.coli binds specifically to a site within the female specific exon of dsx pre-mRNA. This site is required for female specific splicing and female specific polyadenylation. Results suggest that tra2 is a positive regulator of dsx pre-mRNA processing.
                      Cotransfection analyses in which dsx, tra and tra2 cDNAs are expressed in Kc cells revealed that female specific splicing of dsx transcript is positively regulated by tra and tra2 gene products. Analysis of mutant constructs of dsx demonstrates that a portion of the female specific exon is required for regulation of dsx pre-mRNA splicing.
                      Cotransfection assays to examine regulatory interactions between specific cis-acting sequence elements of dsx pre-mRNA, and tra and tra2 gene products establish that tra and tra2 function to activate the use of the female specific exon.
                      The tra2 gene product may function to control sexual differentiation by directly regulating the processing of the dsx pre-mRNA.
                      The mechanism of sex determination in the germ line has been analysed.
                      Mutant individuals are female and male intersexuals.
                      The dsx gene regulates sexual differentiation of somatic tissues. Null alleles convert chromosomally male and female flies into sterile intersexes of similar phenotype. Dominant alleles (e.g., dsxD, dsxM, dsxT) transform females into intersexes when heterozygous with a normal allele, and into phenotypic males when homozygous or heterozygous with a dsx-null allele or deficiency, but they have no effect in males. Most alleles at dsx affect both sexes; however, some alleles affect only one sex. The recessive allele dsx11 converts males into intersexes and is complemented by dominant dsx alleles and recessive alleles that affect only females (dsx22) (Baker and Ridge, 1980; Nothiger et al., 1987). Double-mutant combinations of dsx null mutations with loss-of-function alleles at tra, tra2 and ix result in a doublesex phenotype (Mukherjee and Hildreth, 1971; Baker and Ridge, 1980; Nothiger et al., 1987). Double-mutant combinations of dsxD/+ with null alleles of tra and tra2 convert females into phenotypic males or with ix into more male-like intersexes (Baker and Ridge, 1980; Nothiger et al., 1987). The dose of dsx alleles can alter the phenotype; triploid female flies dsxD/+/+ are sterile and with a weak external dsx phenotype (Gowen and Fung, 1957; Nothiger et al., 1987); diploid female flies that are dsxD/+, but also carry a dsx+ duplication Tp(3;Y)P92 are sterile but female in appearance (Nothiger et al., 1987). Germline sexual differentiation is not dependent on dsx+ function; only the chromosomal constitution determines the sex of transplanted dsxM/+, dsx1, dsxD/+ and dsxD/dsx1 germ cells (Nothiger, Roost and Schupbach, 1980; Schupbach, 1982). The dsx+ gene does not appear to encode any vital functions (Baker and Ridge, 1980). The normal body size differences between male and female flies is maintained in dsx-null mutants (Hildreth, 1965) and in females heterozygous for dsxD/+, dsxD/dsx1 (Fung and Gowen, 1957; Baker and Ridge, 1980) and dsxM/+ (Nothiger et al., 1987). The sexcomb bristles on the prothoracic basitarsus in both sexes of dsx-null homozygotes (Hildreth, 1965; Mukherjee and Hildreth, 1971; Baker and Ridge, 1980) and female dsxD/dsx1 (Nothiger et al., 1987) are intermediate in number, morphology and position compared with the sexcomb bristles in normal males and the transverse row bristles in normal females. The central sexcomb bristle is retained in dsx-null mutants (Hildreth, 1965). In dsx-null mutants, the pigmentation of the fifth tergite is intermediate between the completely pigmented male and the posteriorly pigmented female tergite, whereas the sixth tergite is darkly pigmented (Hildreth, 1965; Baker and Ridge, 1980). Female flies that are dsxD/+ or dsxM/+ are similar to dsx homozygotes (Fung and Gowen, 1957; Duncan and Kaufman, 1975; Baker and Ridge, 1980; Nothiger et al. 1987). Male dsx flies have a seventh tergite and sternite with bristles (Hildreth, 1965; Baker and Ridge, 1980). Female flies heterozygous for dominant alleles and either dsx1 or dsx deficiencies have the male number of tergites and sternites with the male pattern of pigmentation (Duncan and Kaufman, 1975; Baker and Ridge, 1980; Nothiger, Roost and Schupach, 1980; Nothiger et al., 1987). By clonal analysis, the action of dsx has been shown to be cell autonomous in the differentiation of the sexcombs and pigmentation of the abdominal tergites; dsx+ is required until the end of the larval period for the proper sexual differentiation of the sexcombs and into the pupal period, close to the time of the termination of divisions of the abdominal histoblasts, for proper sexual differentiation of the abdominal histoblasts and for proper sexual differentiation of the abdomen (Baker and Ridge, 1980). Both male and female genitalia are formed in dsx null mutant flies and in female flies heterozygous for dominant alleles (Fung and Gowen, 1957; Hildreth, 1965; Epper, 1981; Nothiger et al., 1987); a second penis differentiates with a reduced aedeagus and parameres within the female vaginal area (Hildreth, 1965). In dsxD/+ females, the development of the female genitalia and second penis are very similar to that of dsx-null flies, whereas in dsxM/+ females the female genitalia are more severely reduced (Gowen and Fung, 1975; Baker and Ridge, 1980; Nothiger, Roost and Schupach, 1980; Epper, 1981; Nothiger et al., 1987). Male genitalia from dsx null flies and females heterozygous for dsx dominant alleles contain all elements except a basal apodeme but other external structures such as the penis and accessory elements are reduc
                      Origin and Etymology
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                      Synonyms and Secondary IDs (16)
                      Reported As
                      Symbol Synonym
                      Hr
                      dsx
                      (Aranha and Vasconcelos, 2018, Wu et al., 2018, Chen et al., 2017, Hu et al., 2017.6.13, Mohr et al., 2017, Transgenic RNAi Project members, 2017-, Wagamitsu et al., 2017, Yamamoto and Kohatsu, 2017, Auer and Benton, 2016, Clandinin and Owens, 2016-, Coen and Murthy, 2016, Kalay et al., 2016, Massey and Wittkopp, 2016, Meissner et al., 2016, Pavlou et al., 2016, Regan et al., 2016, Signor et al., 2016, Chatterjee et al., 2015, Fear et al., 2015, Gene Disruption Project members, 2015-, Lucchesi and Kuroda, 2015, Luo and Baker, 2015, Price et al., 2015, Schertel et al., 2015, Verhulst and van de Zande, 2015, Ashwal-Fluss et al., 2014, Atallah et al., 2014, Feng et al., 2014, Ma et al., 2014, Rezával et al., 2014, Saccone et al., 2014, Billeter and Levine, 2013, Castellanos et al., 2013, Crickmore and Vosshall, 2013, Devi and Shyamala, 2013, Fernández and Kravitz, 2013, Haussmann et al., 2013, Park and Burtis, 2013.3.21, Pavlou and Goodwin, 2013, Shirangi et al., 2013, Spokony, 2013.8.25, Yamamoto and Koganezawa, 2013, Foronda et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Ruben et al., 2012, Tarone et al., 2012, Venables et al., 2012, Whitworth et al., 2012, Abruzzi et al., 2011, Bickel et al., 2011, Chang et al., 2011, Chatterjee et al., 2011, Gempe and Beye, 2011, Graham et al., 2011, Graveley et al., 2011, Hartmann et al., 2011, Jungreis et al., 2011, Kimura, 2011, Luo et al., 2011, McNeil et al., 2011, Pan et al., 2011, Sarno et al., 2011, Tanaka et al., 2011, von Philipsborn et al., 2011, Fernández-Ayala et al., 2010, Mellert et al., 2010, Rideout et al., 2010, Robinett et al., 2010, Ruiz and Sanchez, 2010, Alvarez et al., 2009, Ji and Tulin, 2009, Lebo et al., 2009, Ruedi and Hughes, 2009, Schuettengruber et al., 2009, Shen et al., 2009, Song et al., 2009, Stone and Ayroles, 2009, Camara and Doren, 2008, DeFalco et al., 2008, Fujii et al., 2008, Hempel and Oliver, 2008, Kalamegham and Oliver, 2008, Kimura et al., 2008, Saccone et al., 2008, Sanders and Arbeitman, 2008, Sanders and Arbeitman, 2008, Siera and Cline, 2008, Williams et al., 2008, Yang et al., 2008, Chintapalli et al., 2007, Goldman and Arbeitman, 2007, Graze et al., 2007, Hempel and Oliver, 2007, Kitadate et al., 2007, Muse et al., 2007, Nurminsky, 2007, Qi et al., 2007, Rideout et al., 2007, Siera and Cline, 2007, Sofola et al., 2007, Billeter et al., 2006, Casper and Van Doren, 2006, Douglas and Levine, 2006, Le Bras and Van Doren, 2006, Lee et al., 2006, McIntyre et al., 2006, Sciabica and Hertel, 2006, Shigenobu et al., 2006, Shirangi et al., 2006, Zhang et al., 2006, Barmina et al., 2005, Bayrer et al., 2005, Gleason, 2005, Haag and Doty, 2005, Tarone et al., 2005, Wawersik et al., 2005, Yamamoto et al., 2004, Chandler et al., 2003, Lalli et al., 2003, Svensson et al., 2003, Hall, 2002, Lee et al., 2002, An et al., 2000, Amrein et al., 1994)
                      ix-62c
                      Name Synonyms
                      Hermaphrodite
                      double sex
                      intersex-62c
                      Secondary FlyBase IDs
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                        References (537)