General Information
Symbol
Dmel\da
Species
D. melanogaster
Name
daughterless
Annotation Symbol
CG5102
Feature Type
FlyBase ID
FBgn0267821
Gene Model Status
Stock Availability
Gene Snapshot
Daughterless is a class I bHLH protein important for sex determination and dosage compensation by controlling the feminizing switch gene Sxl. It participates in transcriptional regulation of a wide variety processes, including oogenesis, neurogenesis, myogenesis and cell proliferation. [Date last reviewed: 2016-06-30]
Genomic Location
Cytogenetic map
Sequence location
2L:10,388,105..10,393,228 [+]
Recombination map
2-42
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
BASIC HELIX-LOOP-HELIX TRANSCRIPTION FACTORS -
Basic helix-loop-helix (bHLH) transcription factors are sequence-specific DNA-binding proteins that regulate transcription. They are characterized by a 60 amino acid region comprising a basic DNA binding domain followed by a HLH motif formed from two amphipathic α-helices connected by a loop. bHLH transcription factors form homo- and hetero-dimeric complexes, which bind to a E box consensus sequence. (Adapted from PMID:15186484).
UniProt Contributed Function Data
Daughterless/Achaete-scute complex heterodimers act as transcriptional activators of neural cell fates and are involved in sex determination.
(UniProt, P11420)
Phenotypic Description from the Red Book (Lindsley and Zimm 1992)
da: daughterless (C. Cronmiller and T.W. Cline)
da+ performs multiple roles during development. Maternally supplied da+ is required in female embryos as a positive activator of the gene, Sex-lethal (Sxl), the key binary switch gene for the sex determination pathway. Also, da+ expression is required in the somatic gonad of adult females for proper egg membrane formation, and hence for the survival of all progeny regardless of their sex. Embryonic expression of da+ is required in both sexes for the formation of the peripheral nervous system (PNS) and parts of the central nervous system (CNS). And, during larval and/or pupal stages, da+ may be required for the growth and/or differentiation of cells that form the adult cuticle. Amorphic alleles (da2, da3, da5, etc.) are recessive lethals, with a lethal period which is predominantly embryonic (Cronmiller and Cline, 1987; Caudy et al., 1988a). In addition, the hypomorphic allele, da1 (originally called da), is hemizygous [da1/Df(2L)da-] lethal (Mange and Sandler, 1973), and da1 homozygotes die when they undergo the first half of embryonic development at 29 (Cline, 1976). Death appears to be a consequence of dosage compensation defects (Cline, 1983a; Gergen, 1987). Viability of da1 homozygotes is improved by the presence of extra X or Y heterochromatin in either the parental female or her progeny (Sandler, 1972; Mason, 1973). Temperature-sensitive lethality of the da1 zygotic lethal effect is not affected by the Sxl genotype (Cline, 1980). da+ is not required in the germline, since da-(da2/da3) pole cells produce fertile gametes; however, mitotic recombination failed to yield significant da-(da2 or da3) somatic clones, suggesting da+ may be essential during epidermal development (Cronmiller and Cline, 1987). Embryos, homozygous for lethal da alleles, have a reduced CNS, lack all peripheral neurons, and have no external sensory structures (Caudy et al., 1988a). Adult flies heterozygous for a deletion of the achaete-scute (ASC) genes and simultaneously heterozygous for Df(da) (also da2/+ or da5/+) exhibit characteristic bristle defects (Dambly-Chaudiere et al., 1988). Hemizygosity for da+ reduces the number of supernumerary bristles in Hw mutants (Dambly-Chaudiere et al., 1988). In addition to its zygotic phenotype, da1 exhibits two separable maternal effects. There is a female-specific maternal effect: At 22 and 25, homozygous da1 females produce no daughters, while at 18, they produce approximately 20% as many daughters as sons (Cline, 1976). At 29, da1 displays a sex-nonspecific maternal effect. Homozygous females are reversibly sterile; they lay eggs that show little or no development (Cline, 1976). Sterility of da1 females at high temperature results from a defect in the somatic gonad rather than in the germline, since da- germ cells in wild-type ovaries produce normal eggs which support full viability of sons (Cronmiller and Cline, 1987). The female-specific maternal effect has a temperature-sensitive period which includes the last 60 hr of oogenesis and the first 3 hr of development (Cline, 1976). This maternal effect is also observed in crosses of da1 females to D. simulans males (Watanabe and Yamada, 1977). The female-lethal maternal effect is autonomous to the germline, as demonstrated by transplantation of da1, or da2/da3 pole cells into + hosts (Cline, 1983b; Cronmiller and Cline, 1987). Female zygotes from da1 mothers at 25 die as embryos. Such lethal female embryos show consistent abnormalities in midgut formation, and in about 50% of the abnormal embryos, shortening of the germ band fails, while anus and posterior spiracles open on the dorsal surface behind the head segments (Counce). Female embryos from da1 mothers also show consistent defects in the CNS, which is either reduced in width or shows abrupt bends or twists; abnormally formed gut often extends into the CNS (Caudy et al., 1988a). The majority of daughters of da1 mothers surviving at 18 are morphologically abnormal, often missing structures from one or more imaginal discs or abdominal histoblasts, and frequently with duplication of structures (Cline, 1976). Though it was reported that daughters of homozygous da1 females could be rescued by cytoplasmic injection (Bownes et al., 1977), the apparent rescue was subsequently found to result from nonspecific effects that may have slowed the early development of females who are on the threshold of surviving (Cline, 1984; see also Muir and Bell, 1987). Gynandromorphs can survive the lethal maternal effects, but there is no localized lethal focus. Diplo-X tissue develops abnormally alongside normally developing haplo-X tissue. Survival of the mosaics and their average fraction of diplo-X tissue increases with decreasing temperature (Cline, 1976). The da1 maternal effect masculinizes escaper daughters that are homozygous for mle (Cline, 1984) and masculinizes triploid intersex (XXAAA) progeny (Cline, 1983a). Females heterozygous for Sxl alleles that lead to male development develop as sterile males, mosaic intersexes, or sterile females (depending on the Sxl allele), when produced by da1 mothers (Cline, 1984). The da1 female lethal maternal effect is unaffected by tra or dsx (Bell, 1954; Colainne and Bell, 1968). However, daughters of da1/da1 mothers are almost fully rescued by a single zygotic dose of SxlM1 and to a limited degree by a duplication for Sxl+ (Cline, 1978). Conversely, zygotic Sxl- enhances the da maternal effect. Females with reduced Sxl dose (Sxl-/+) fail to survive from da1/da1 mothers at the semipermissive 18 (Cline, 1978). A strong dominant da maternal effect [da1/+, Df(2L)da/+, or da2/+ mothers] is observed when female progeny are doubly heterozygous for Sxl- and sis-a- (Cline, 1986, Genetics 113: 641-63; Cronmiller and Cline, 1986, 1987). The maternal effect of da1 is made semidominant also by E(da) (cis or trans to da1) in the mother (Mange and Sandler, 1973; see also Cline, 1980). The zygotic da+ dose itself does not affect expression of Sxl+ sex determination function (Cronmiller and Cline, 1986).
Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
2

Please see the GBrowse view of Dmel\da or the JBrowse view of Dmel\da 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
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.45
gene_with_stop_codon_read_through ; SO:0000697
Stop-codon suppression (UAG) postulated; FBrf0234051.
Gene model reviewed during 6.25
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0080008
3162
710
FBtr0332195
3603
710
FBtr0332196
3103
710
FBtr0474232
3162
775
Additional Transcript Data and Comments
Reported size (kB)
3.4, 3.2, 2.954 (longest cDNA)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0079598
73.9
710
7.08
FBpp0304504
73.9
710
7.08
FBpp0304505
73.9
710
7.08
FBpp0423185
81.3
775
7.26
Polypeptides with Identical Sequences

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

710 aa isoforms: da-PA, da-PB, da-PC
Additional Polypeptide Data and Comments
Reported size (kDa)
82 (kD observed); 74 (kD predicted)
710 (aa); 74 (kD predicted)
Comments
External Data
Subunit Structure (UniProtKB)
Homodimer. Efficient DNA binding requires dimerization with another bHLH protein. Interacts with Amos. Interacts (via bHLH motif) with sisA and sc. Interacts with dpn (via bHLH motif).
(UniProt, P11420)
Crossreferences
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\da 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 (26 terms)
Molecular Function (11 terms)
Terms Based on Experimental Evidence (9 terms)
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:ac; FB:FBgn0000022
inferred from physical interaction with FLYBASE:amos; FB:FBgn0003270
inferred from physical interaction with FLYBASE:ase; FB:FBgn0000137
inferred from physical interaction with FLYBASE:HLH54F; FB:FBgn0022740
inferred from physical interaction with FLYBASE:emc; FB:FBgn0000575
inferred from physical interaction with FLYBASE:ato; FB:FBgn0010433
inferred from physical interaction with UniProtKB:P10627
(assigned by UniProt )
inferred from physical interaction with FLYBASE:ase; FB:FBgn0000137
inferred from physical interaction with FLYBASE:l(1)sc; FB:FBgn0002561
inferred from physical interaction with FLYBASE:sc; FB:FBgn0004170
inferred from physical interaction with FLYBASE:ey; FB:FBgn0005558
inferred from genetic interaction with FLYBASE:E(spl)m7-HLH; FB:FBgn0002633
inferred from genetic interaction with FLYBASE:E(spl)mγ-HLH; FB:FBgn0002735
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000927455
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000927455
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000927455
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000927455
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000927455
(assigned by GO_Central )
Biological Process (13 terms)
Terms Based on Experimental Evidence (11 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000927455
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000927455
(assigned by GO_Central )
Cellular Component (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000927455
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000927455
(assigned by GO_Central )
Expression Data
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: maternally deposited

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
There are two phases of da expression during oogenesis. In the first phase, da transcripts are detected throughout the germarium and in all egg chambers in both germ line and the somatic cells until approximately stage S3. After stage S3, transcripts are not detected in the germ line and are detected at low levels in the follicle cells. The late expression phase starts in stage S8 at which point strong expression is observed in the germ line in the nurse cells and increases until late stage 10. At that point transfer of da transcripts to the oocyte is initiated. Weak expression continues to be observed in follicle cells.
Both da transcripts are present at constant proportion in all stages of development. The 3.2 kb transcript is slightly overrepresented in 0-2.5 hr embryos (in comparison to other developmental stages).
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
da is expressed at a low level in ato positive cells in the morphogenetic furrow in third instar larval eye discs. It is expressed at a high level in non-neuronal cells surrounding ato-positive R8 cells between proneural clusters posterior to the morphogenetic furrow.
da protein expression is first seen as a dorsoventral stripe, several cells wide, at the anterior side of the morphogenetic furrow that quickly resolves into expression in single cells within the 2-3 ommatidial rows in the posterior portion of the furrow. The single da-expressing cells appear to be R8 cells. da protein expression exactly corresponds with ato protein expression.
da protein is widely distributed throughout the somatic component of the ovary. Expression is first observed in prefollicular cells and follicle cells from germarium region 2 until stage S3. The most intense staining is seen at the anterior portion of the germarium at stage 3 as a nascent follicle is ready to pinch off from the germarium. In egg chambers through stage S3, da protein expression is maintained in all the nuclei of follicular epithelial cells. Protein levels diminish in the follicular epithelium thereafter. Throughout egg chamber maturation, expression is observed in interfollicular stalk cells and interfollicular polar cells. In stage S9 and beyond, uniform levels of da protein are observed in all follicle cells.
da protein is observed in the nuclei of most cells during embryogenesis but is not present in pole cells. da protein is present throughout the preblastoderm embryo and disappears before blastoderm formation. Protein levels increase again before germ band extension and reach maximal levels during stages 9-11. da protein is present in ectodermal cells as well as in putative neuroblasts during the process of neuroblast delamination and after. The levels of da protein are fairly uniform across the ectodermal layer. Neuronal precursors, however, appear to have elevated levels of da protein. da protein expression is reduced in most tissue types during germ band retraction. At later stages higher levels of da protein are seen n particular tissues including a subset of CNS cells, salivary glands, and parts of the gut and muscles. In wing imagingal discs, a uniform level of da protein is observed in all epidermal cells. An elevated protein level is observed in some neuronal precursor cells. In leg discs, elevated da protein levels are seen in the large cluster of SOPS that will later form the chordotonal organ. In eye discs, elevated da protein levels are seen in cells posterior to the morphogenetic furrow that are thought to correspond to R8 cells.
da protein is detected at all developmental stages on western blots. Peak levels are observed in 5-12hr embryos. da protein was found to be expressed continuously and ubiquitously during embryogenesis by immunolocalization. In later stages, protein levels are highest in the supraoesophageal ganglion, the ventral cord and the salivary gland. The only nuclei that appear not to contain da protein are the vitellophages. In larvae, the highest levels of da protein are observed in the imaginal discs (eye-antenna, wing, leg, haltere, and labial), in salivary glands, and in a subset of cells in the CNS. During oogenesis, da protein is detected in follicle cells but not in the germline. Finally, da protein is detected in male gona s. In the testis, light staining is observed in apically located cells and heavy staining is observed in somatic cyst cells and in terminal epithelial cells. Intense staining is also observed in the seminal vescicle and the anterior ejaculatory duct epithelia.
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\da 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
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Images
Alleles, Insertions, Transgenic Constructs and Phenotypes
Classical and Insertion Alleles ( 41 )
For All Classical and Insertion Alleles Show
 
Allele of da
Class
Mutagen
Associated Insertion
Stocks
Known lesion
    0
    Yes
      0
      --
      Other relevant insertions
      miscellaneous insertions
      Name
      Expression Data
      Transgenic Constructs ( 29 )
      For All Alleles Carried on Transgenic Constructs Show
      Transgenic constructs containing/affecting coding region of da
      Allele of da
      Mutagen
      Associated Transgenic Construct
      Stocks
      Transgenic constructs containing regulatory region of da
      GAL4 construct
      vital-reporter construct
      Name
      Expression Data
      Deletions and Duplications ( 18 )
      Summary of Phenotypes
      For more details about a specific phenotype click on the relevant allele symbol.
      Lethality
      Allele
      lethal (with da1)
      lethal (with da7)
      Sterility
      Allele
      Other Phenotypes
      Allele
      Phenotype manifest in
      Allele
      Orthologs
      Human Orthologs (via DIOPT v7.1)
      Homo sapiens (Human) (3)
      Species\Gene Symbol
      Score
      Best Score
      Best Reverse Score
      Alignment
      Complementation?
      Transgene?
      11 of 15
      Yes
      Yes
      11 of 15
      Yes
      Yes
       
       
      11 of 15
      Yes
      Yes
      Model Organism Orthologs (via DIOPT v7.1)
      Mus musculus (laboratory mouse) (3)
      Species\Gene Symbol
      Score
      Best Score
      Best Reverse Score
      Alignment
      Complementation?
      Transgene?
      10 of 15
      Yes
      Yes
      9 of 15
      No
      Yes
      9 of 15
      No
      Yes
      Rattus norvegicus (Norway rat) (3)
      9 of 13
      Yes
      Yes
      7 of 13
      No
      Yes
      7 of 13
      No
      Yes
      Xenopus tropicalis (Western clawed frog) (3)
      5 of 12
      Yes
      Yes
      5 of 12
      Yes
      Yes
      3 of 12
      No
      Yes
      Danio rerio (Zebrafish) (4)
      11 of 15
      Yes
      Yes
      11 of 15
      Yes
      Yes
      8 of 15
      No
      Yes
      3 of 15
      No
      Yes
      Caenorhabditis elegans (Nematode, roundworm) (1)
      8 of 15
      Yes
      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) ( EOG091906QL )
      Organism
      Common Name
      Gene
      AAA Syntenic Ortholog
      Multiple Dmel Genes in this Orthologous Group
      Drosophila melanogaster
      fruit fly
      Drosophila suzukii
      Spotted wing Drosophila
      Drosophila simulans
      Drosophila sechellia
      Drosophila erecta
      Drosophila yakuba
      Drosophila ananassae
      Drosophila pseudoobscura pseudoobscura
      Drosophila persimilis
      Drosophila willistoni
      Drosophila virilis
      Drosophila mojavensis
      Drosophila grimshawi
      Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091505GW )
      Organism
      Common Name
      Gene
      Multiple Dmel Genes in this Orthologous Group
      Musca domestica
      House fly
      Glossina morsitans
      Tsetse fly
      Lucilia cuprina
      Australian sheep blowfly
      Lucilia cuprina
      Australian sheep blowfly
      Mayetiola destructor
      Hessian fly
      Mayetiola destructor
      Hessian fly
      Aedes aegypti
      Yellow fever mosquito
      Anopheles darlingi
      American malaria mosquito
      Anopheles gambiae
      Malaria mosquito
      Culex quinquefasciatus
      Southern house mosquito
      Culex quinquefasciatus
      Southern house mosquito
      Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0BE2 )
      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 florea
      Little honeybee
      Apis mellifera
      Western honey bee
      Bombus impatiens
      Common eastern bumble bee
      Bombus terrestris
      Buff-tailed bumblebee
      Linepithema humile
      Argentine ant
      Megachile rotundata
      Alfalfa leafcutting bee
      Nasonia vitripennis
      Parasitic wasp
      Dendroctonus ponderosae
      Mountain pine beetle
      Tribolium castaneum
      Red flour beetle
      Pediculus humanus
      Human body louse
      Cimex lectularius
      Bed bug
      Acyrthosiphon pisum
      Pea aphid
      Zootermopsis nevadensis
      Nevada dampwood termite
      Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0BBN )
      Organism
      Common Name
      Gene
      Multiple Dmel Genes in this Orthologous Group
      Strigamia maritima
      European centipede
      Stegodyphus mimosarum
      African social velvet spider
      Stegodyphus mimosarum
      African social velvet spider
      Tetranychus urticae
      Two-spotted spider mite
      Daphnia pulex
      Water flea
      Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0NNT )
      Organism
      Common Name
      Gene
      Multiple Dmel Genes in this Orthologous Group
      Strongylocentrotus purpuratus
      Purple sea urchin
      Human Disease Model Data
      FlyBase Human Disease Model Reports
      Alleles Reported to Model Human Disease (Disease Ontology)
      Download
      Models ( 5 )
      Allele
      Disease
      Evidence
      References
      inferred from mutant phenotype
      inferred from mutant phenotype
      inferred from mutant phenotype
      inferred from mutant phenotype
      inferred from mutant phenotype
      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.
      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.
      Dmel gene
      Ortholog showing functional complementation
      Supporting References
      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
      Summary of Genetic Interactions
      esyN Network Diagram
      esyN Network Key:
      Suppression
      Enhancement

      Please look at the allele data for full details of the genetic interactions
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      External Data
      Subunit Structure (UniProtKB)
      Homodimer. Efficient DNA binding requires dimerization with another bHLH protein. Interacts with Amos. Interacts (via bHLH motif) with sisA and sc. Interacts with dpn (via bHLH motif).
      (UniProt, P11420 )
      Linkouts
      Pathways
      Gene Group - Pathway Membership (FlyBase)
      External Data
      Linkouts
      Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
      Genomic Location and Detailed Mapping Data
      Chromosome (arm)
      2L
      Recombination map
      2-42
      Cytogenetic map
      Sequence location
      2L:10,388,105..10,393,228 [+]
      FlyBase Computed Cytological Location
      Cytogenetic map
      Evidence for location
      31D11-31E1
      Limits computationally determined from genome sequence between P{lacW}RnrLk06709 and P{lacW}KdelRk00311
      Experimentally Determined Cytological Location
      Cytogenetic map
      Notes
      References
      31E-31E
      (determined by in situ hybridisation)
      Experimentally Determined Recombination Data
      Location
      2-39.3
      2-41.3
      Left of (cM)
      Right of (cM)
      Notes
      Stocks and Reagents
      Stocks (24)
      Genomic Clones (19)
      cDNA Clones (81)
       

      Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see GBrowse for alignment of the cDNAs and ESTs to the gene model.

      cDNA clones, fully sequences
      BDGP DGC clones
      Other clones
      Drosophila Genomics Resource Center cDNA clones

      For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

      cDNA Clones, End Sequenced (ESTs)
      RNAi and Array Information
      Linkouts
      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: da l(2)k08611
      Additional comments
      Other Comments
      DNA-protein interactions: genome-wide binding profile assayed for da protein in 2-3 hr embryos; see BDTNP1_TFBS_da collection report.
      da has a role in somatic cell proliferation during oogenesis. da is required for complete differentiation of polar and stalk follicle cells.
      Analysis of the function of da.
      Dominant interactions indicate that toc is acting in the same signalling pathways for the formation of the egg chamber as da, N and Dl.
      Four genes whose products are required for various stages of the cell cycle are misexpressed in the PNS of da mutant embryos.
      da is autonomously required for the neurogenesis of all photoreceptor cells that differentiate within the furrow. Both da and ato are independently activated within the eye disc, proper maintenance of both da and ato expression is dependent on the other protein. Loss of da disrupts the progression of the morphogenetic furrow and this effect is mediated by the loss of both hh and dpp. Also da function is necessary for reentry into the cell cycle by cells of the second mitotic wave posterior to the morphogenetic furrow.
      Proneural and neurogenic genes control specification and morphogenesis of stomatogastric nerve cell precursors.
      E(spl)-complex bHLH proteins interact with proneural proteins, with members of the E(spl) family exhibiting distinct preferences for different proneural proteins.
      Immunoprecipitation experiments suggest that sc and da form a heteromeric complex in vivo.
      The da gene product exerts a regulatory effect on Sxl expression.
      Overexpression of da using the GAL4 system, but not the ectopic expression of the AS-C genes l(1)sc or sc, leads to the formation of ectopic neural cells in embryonic tissue without neural competence. This effect os strongly enhanced by coexpressing l(1)sc or sc. Expression of da and/or l(1)sc is not sufficient to overcome the lateral inhibition in the analgen of the embryonic nervous system.
      Transfection assays and in vitro DNA binding experiments indicate that da/sc heterodimers directly activate the Sxl early promoter by binding to both high and low affinity sites. dpn protein represses this activation by specific binding to a unique site within the Sxl early promoter.
      fs(1)Yb is required in the soma for ovary follicle cell differentiation and to support later stages of egg maturation. Mutations at fs(1)Yb show genetic interactions with the N group of neurogenic genes.
      The yeast two hybrid system has been used to demonstrate specific interactions within the sisA, sc, dpn and da group of gene products, and to delimit their interaction domains. The results support and extend the model of the molecular basis of the X/A ratio signal.
      da is required for the survival of salivary gland cells after they invaginate.
      emc forms heterodimers with the ac, sc, l(1)sc, and da products. emc inhibits DNA-binding of ac, sc and l(1)sc/da heterodimers and da homodimers.
      Proneural gene products (ac, da and l(1)sc) activate transcription of Dl in the neuroectoderm by binding to specific sites within its promoter. This transcriptional activation enhances lateral inhibition and helps ensure that cells in the vicinity of prospective neuroblasts will themselves become epidermoblasts.
      DNaseI footprinting analysis of bacterially expressed da and l(1)sc demonstrates that the gene products can bind as heterodimers to different E-box sequence upstream of the ac gene.
      A fragment of the D.melanogaster da gene has been used as a probe for in situ hybridisation of Chrysomya rufifacies polytene chromosomes.
      da is not required for the initial appearance of nascent neuronal precursors but is required for these cells to express multiple neuronal precursor genes and to produce the normal number of neurons.
      Functional analysis of Brer\E12 performed using Drosophila as an in vivo system demonstrates the HLH domain of Brer\E12 can carry out most of the functions performed by the corresponding region of da.
      Ten (unnamed) recessive lethal alleles have been isolated during a cytogenetic analysis of chromosomal region 31.
      da encodes a nuclear protein that is widely expressed throughout development.
      dl gene product interacts with members of the HLH family, including da, ac and sc and dosage sensitive interactions that exist between dl, da, ac, sc and twi are required for the specification of both the embryonic mesoderm and neuroectoderm.
      Extra copies of da+ do not effect male lethality of schb.PP.
      dpn expression in da mutant embryos has been examined: all neural precursors are abnormal, do not express dpn and produce very few neurons. Later in development dpn is found in some CNS neurons.
      Mutations at da, hup and dal loci have no effect on position-effect variegation. Mutant alleles are sensitive to amounts of heterochromatin in the genome.
      There is a weak female-specific dominant synergism between run and da mutations.
      In cotransfection studies the highest levels of ac expression are achieved when a combination of ac and da or sc expression vectors are present in the cotransfection mixture.
      In vitro DNA binding assays using gel retardation to an ac promoter region and hb zygotic promoter region target sequence demonstrates that da protein elicits a weak homodimeric binding and da/ac or da/sc heterodimers bind tightly. Single copy yeast promoters under the control of the GAL4 promoter were used to test whether ac, sc and da proteins could activate transcription of a Ecol\lacZ reporter gene in the yeast assay system. da produces slight activation and the presence of da gives strong activation of the reporter gene. Results suggest that da/l(1)sc heterodimers can function as transcriptional activators in direct proportion to their DNA-binding affinities.
      sisA gene product acts in combination with maternally encoded products of signal transduction genes, da, which communicate the number of X chromosomes to Sxl and thereby determine sexual fate.
      da is required in all regions of the embryo to activate Sxl.
      Mutations in maternal class gene da interact with RpII140wimp.
      DNA sequence analysis reveals four E box binding sites, for the binding of hetero-oligomeric complexes composed of da or AS-C proteins, in the first 877 bp of the ac upstream region. Electrophoretic mobility shift assays demonstrate that the emc protein can specifically antagonise DNA binding of the da/AS-C complexes in vitro in a dose-dependent manner, h and E(spl) proteins fail to exhibit this inhibitory effect.
      da alleles act as enhancers of spl alleles of N.
      The da gene product may be capable of functionally complementing the genes of ASC.
      Extra wild type copies of da are not sufficient to rescue the female lethality of hhb.PP. da and sc are both required for the induction of Sxl expression.
      E12 and E47 binding factors bind to the murine immunoglobulin kappa chain enhancer. These cDNAs show sequence similarity with da.
      Defects in neural development caused by mutations in da can be prevented by the presence of mutations in N, bib, mam, neu, Dl amd E(spl).
      The lack of interaction between Tpl and Sxl or da suggests that Tpl does not function in measuring the X/A ratio in Drosophila.
      Double hemizygotes for the achaete-scute complex and da <up>Df(1)260-1/+; Df(2L)J27/+</up> or the achaete-scute complex and Df(4)M101-62f show loss of macrochaetae, which none of the single hemizygotes does.
      Duplications carrying wild type and mutant da alleles have been used to determine the relationship between maternal and zygotic activities of the da locus and whether zygotic functions of da are important in the control of Sxl sex determination activities. Individuals with three doses of da+ show no decrease in viability or fertility. Increasing the maternal and zygotic da+ dose reduces the severity of the recessive lethal effect of mutant da alleles and reducing the maternal da+ dose reduces female lethal maternal effect of mutant da alleles. da+ zygotic function is not involved in Sxl+ regulation: lowered da+ zygotic dose does not reduce Sxl+ expression of sex determination functions.
      The interaction between Sxl and da in triploids has been studied.
      da+ performs multiple roles during development. Maternally supplied da+ is required in female embryos as a positive activator of the gene, Sex-lethal (Sxl), the key binary switch gene for the sex determination pathway. Also, da+ expression is required in the somatic gonad of adult females for proper egg membrane formation and hence for the survival of all progeny regardless of their sex. Embryonic expression of da+ is required in both sexes for the formation of the peripheral nervous system (PNS) and parts of the central nervous system (CNS). And, during larval and/or pupal stages, da+ may be required for the growth and/or differentiation of cells that form the adult cuticle. Amorphic alleles (da2, da3, da5, etc.) are recessive lethals, with a lethal period which is predominantly embryonic (Cronmiller and Cline, 1987; Caudy et al., 1988). In addition, the hypomorphic allele, da1 (originally called da), is hemizygous (da1/da deficiencies) lethal (Mange and Sandler, 1973), and da1 homozygotes die when they undergo the first half of embryonic development at 29oC (Cline, 1976). Death appears to be a consequence of dosage compensation defects (Cline, 1983a; Gergen, 1987). Viability of da1 homozygotes is improved by the presence of extra X or Y heterochromatin in either the parental female or her progeny (Sandler, 1972; Mason, 1973). Temperature-sensitive lethality of the da1 zygotic lethal effect is not affected by the Sxl genotype (Cline, 1980). da+ is not required in the germ-line, since da- (da2/da3) pole cells produce fertile gametes; however, mitotic recombination failed to yield significant da- (da2 or da3) somatic clones, suggesting da+ may be essential during epidermal development (Cronmiller and Cline, 1987). Embryos, homozygous for lethal da alleles, have a reduced CNS, lack all peripheral neurons and have no external sensory structures (Caudy et al., 1988). Adult flies heterozygous for a deletion of the achaete-scute (ASC) genes and simultaneously heterozygous for Df(da) (also da2/+ or da5/+) exhibit characteristic bristle defects (Dambly-Chaudiere, Ghysen, Jan and Jan 1988). Hemizygosity for da+ reduces the number of supernumerary bristles in Hw mutants (Dambly-Chaudiere, Ghysen, Jan and Jan, 1988). In addition to its zygotic phenotype, da1 exhibits two separable maternal effects. There is a female-specific maternal effect: At 22oC and 25oC, homozygous da1 females produce no daughters, while at 18oC, they produce approximately 20% as many daughters as sons (Cline, 1976). At 29oC, da1 displays a sex-nonspecific maternal effect. Homozygous females are reversibly sterile; they lay eggs that show little or no development (Cline, 1976). Sterility of da1 females at high temperature results from a defect in the somatic gonad rather than in the germ-line, since da- germ cells in wild-type ovaries produce normal eggs which support full viability of sons (Cronmiller and Cline, 1987). The female-specific maternal effect has a temperature-sensitive period which includes the last 60 hr of oogenesis and the first 3 hr of development (Cline, 1976). This maternal effect is also observed in crosses of da1 females to D.simulans males (Watanabe and Yamada, 1977). The female-lethal maternal effect is autonomous to the germ-line, as demonstrated by transplantation of da1, or da2/da3 pole cells into + hosts (Cline, 1983b; Cronmiller and Cline, 1987). Female zygotes from da1 mothers at 25oC die as embryos. Such lethal female embryos show consistent abnormalities in midgut formation, and in about 50% of the abnormal embryos, shortening of the germ band fails, while anus and posterior spiracles open on the dorsal surface behind the head segments (Counce). Female embryos from da1 mothers also show consistent defects in the CNS, which is either reduced in width or shows abrupt bends or twists; abnormally formed gut often extends into the CNS (Caudy et al., 1988). The majority of daughters of da1 mothers surviving at 18oC are morphologically abnormal, often missing structures from one or more imaginal discs or abdominal histoblasts and frequently with duplication of structures (Cline, 1976). Though it was reported that daughters of homozygous da1 females could be rescu
      Origin and Etymology
      Discoverer
      Bell.
      Etymology
      Identification
      External Crossreferences and Linkouts ( 58 )
      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/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
      UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
      Other crossreferences
      BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
      Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
      Linkouts
      Drosophila Genomics Resource Center - Drosophila Genomics Resource Center 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
      GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
      iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
      Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
      KEGG Genes - Molecular building blocks of life in the genomic space.
      Synonyms and Secondary IDs (13)
      Reported As
      Symbol Synonym
      l(2)k08611
      Secondary FlyBase IDs
      • FBgn0000413
      • FBgn0022036
      Datasets (2)
      Study focus (2)
      Experimental Role
      Project
      Project Type
      Title
      • bait_protein
      ChIP characterization of transcription factor genome binding, Berkeley Drosophila Transcription Factor Network Project.
      • bait_protein
      Genome-wide localization of transcription factors by ChIP-chip and ChIP-Seq.
      References (479)