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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 (da) encodes 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: 2019-03-07]
Key Links
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.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Summaries
Gene Group (FlyBase)
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).
Protein Function (UniProtKB)
Daughterless/Achaete-scute complex heterodimers act as transcriptional activators of neural cell fates and are involved in sex determination.
(UniProt, P11420)
Phenotypic Description (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).
Summary (Interactive Fly)
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 (27 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 (14 terms)
Terms Based on Experimental Evidence (12 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:sens; FB:FBgn0002573
inferred from genetic interaction with FLYBASE:sc; FB:FBgn0004170
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
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: maternally deposited

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 in cells of the intestinal stem cell lineage in adult posterior midgut.
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, and Transgenic Constructs
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 records found.
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( 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
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (0)
No records found.
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 5 )
Potential Models Based on Orthology ( 3 )
Modifiers Based on Experimental Evidence ( 0 )
Allele
Disease
Interaction
References
Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
 
Disease Associations of Human Orthologs (via DIOPT v7.1 and OMIM)
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 see the Physical Interaction reports below for full details
protein-protein
Physical Interaction
Assay
References
Summary of Genetic Interactions
esyN Network Diagram
esyN Network Key:
Suppression
Enhancement

Please look at the allele data for full details of the genetic interactions
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
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
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Gene Group - Pathway Membership (FlyBase)
External Data
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
31E1-31E1
Left limit from in situ hybridisation (FBrf0067338) Right limit from in situ hybridisation (FBrf0067338); 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 (25)
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 ( 65 )
Sequence Crossreferences
NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
UniProt/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
Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
KEGG Genes - Molecular building blocks of life in the genomic space.
Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
Linkouts
Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
Synonyms and Secondary IDs (13)
Reported As
Symbol Synonym
da
(Ishibashi et al., 2019, Shokri et al., 2019, Bischof et al., 2018, Guo et al., 2018, Knuckles et al., 2018, Lan et al., 2018, Wang and Baker, 2018, Bhattacharya et al., 2017, Kan et al., 2017, Liu and Jin, 2017, Stessman et al., 2017, D'Rozario et al., 2016, Jungreis et al., 2016, Golubyatnikov et al., 2015, Schertel et al., 2015, Tamberg et al., 2015, Chen et al., 2014, Hsiao et al., 2014, Jiang and Singh, 2014, Rogers et al., 2014, Tanaka-Matakatsu et al., 2014, Victorsen and White, 2014.4.15, Bergwitz et al., 2013, Das et al., 2013, Oortveld et al., 2013, Tögel et al., 2013, Distefano et al., 2012, Kvon et al., 2012, Powell et al., 2012, Barry et al., 2011, Bhattacharya and Baker, 2011, Debat et al., 2011, Grice and Liu, 2011, Kuzin et al., 2011, Li et al., 2011, Nègre et al., 2011, Neumüller et al., 2011, Park et al., 2011, Pruteanu-Malinici et al., 2011, Toku et al., 2011, Bardin et al., 2010, Bernard et al., 2010, Ismat et al., 2010, Johnson et al., 2010, Salzer et al., 2010, The modENCODE Consortium, 2010, The modENCODE Consortium, 2010, Aerts et al., 2009, Ozdowski et al., 2009, Schaaf et al., 2009, González et al., 2008, Guntur and Lundell, 2008, Kaspar et al., 2008, Lim et al., 2008, Melicharek et al., 2008, Park et al., 2008, Sukhanova and Du, 2008, Vonkavaara et al., 2008, Wong et al., 2008, Yasugi et al., 2008, Christensen and Cook, 2007.10.29, Christensen and Cook, 2006.8.30, Hossain et al., 2005, Macdonald and Long, 2005, Schlatter and Maier, 2005, Sawamura and Yamamoto, 2004, Castanon et al., 2001, Jarman et al., 1993)
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 (491)