FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\bs
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General Information
Symbol
Dmel\bs
Species
D. melanogaster
Name
blistered
Annotation Symbol
CG3411
Feature Type
FlyBase ID
FBgn0004101
Gene Model Status
Stock Availability
Gene Summary
blistered (bs) encodes the Drosophila ortholog of mammalian Serum Response Factor (SRF). It is a sequence-specific transcription factor contributing to intervein cell specification in the wing, tracheal terminal branching and flight muscle maturation. [Date last reviewed: 2021-07-29] (FlyBase Gene Snapshot)
Also Known As

DSRF, SRF, pruned, D-SRF, serum response factor

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
2-106
RefSeq locus
NT_033778 REGION:24342861..24376473
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (21 terms)
Molecular Function (7 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
Biological Process (13 terms)
Terms Based on Experimental Evidence (13 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from expression pattern
involved_in pupal development
inferred from mutant phenotype
involved_in short-term memory
inferred from mutant phenotype
involved_in sleep
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
located_in nucleus
inferred from high throughput direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN004327372
Gene Group (FlyBase)
Protein Family (UniProt)
-
Summaries
Gene Snapshot
blistered (bs) encodes the Drosophila ortholog of mammalian Serum Response Factor (SRF). It is a sequence-specific transcription factor contributing to intervein cell specification in the wing, tracheal terminal branching and flight muscle maturation. [Date last reviewed: 2021-07-29]
Gene Group (FlyBase)
MADS-BOX TRANSCRIPTION FACTORS -
The MADS box transcription factors are sequence-specific DNA binding proteins that regulate transcription. They have a conserved MADS-box motif. The N-terminal half of the motif has the DNA-binding specificity while the C-terminal half required for dimerisation. MADS-box proteins specifically recruit other transcription factors into multi-component regulatory complexes. (Adapted from PMID:7744019).
Protein Function (UniProtKB)
Required for the formation of intervein tissue of the wing. Acts in a dosage-dependent manner to suppress wing vein formation and promote development of intervein cells. Might play a role in the proper formation and maintenance of the trachea.
(UniProt, Q24535)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
bs: blistered
thumb
bs2: blistered-2
Edith M. Wallace, unpublished.
Wings blistered, small, pointed; venation thick and plexus-like with branches from and parallel to L5 beyond second crossvein, where there is a semidominant free vein effect. Temperature sensitive. RK2 at 19; RK3 at 25.
ba: balloon
thumb
ba: balloon
From Bridges and Morgan, 1919, Carnegie Inst. Washington Publ. No. 278: 148.
Wings at first inflated with hemolymph to produce blisters and vesicles; venation weak, plexus like; wings smaller, warped, discolored, and divergent. Effect caused by inadequate contraction of epithelium after inflated state of pupal wing [Waddington, 1940, J. Genet. 41: 75-139 (fig.)]. Sensitive to temperature. RK3 above 25; RK2 at 19 or below.
Summary (Interactive Fly)

transcription factor - MADS-box motif - required for vein/intervein formation in the fly wing - implicated in longer forms of memory formation like synaptic long-term potentiation and depression - required for a working memory that lasts only for a few seconds - acts as a boosting mechanism to sustain FGF-induced terminal branching in the tracheal system.

Gene Model and Products
Number of Transcripts
3
Number of Unique Polypeptides
2

Please see the JBrowse view of Dmel\bs for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q24535)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Gene model reviewed during 5.42

Gene model reviewed during 5.52

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0072271
2221
449
FBtr0290089
3792
355
FBtr0343314
3062
449
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0072178
47.6
449
7.06
FBpp0288528
38.7
355
9.33
FBpp0309973
47.6
449
7.06
Polypeptides with Identical Sequences

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

449 aa isoforms: bs-PA, bs-PC
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\bs using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

-0.94

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

Comment: maternally deposited

Additional Descriptive Data

bs transcripts are expressed ubiquitously in unfertilized eggs. The RNA fades away during germ band extension and zygotic expression is not detected until germ band retraction. At stage 14, expression is observed just beneath the epidermis in a cluster of 6-9 cells in the most anterior portion of each hemisegment from T2 to A8. Expression is also observed in the head, in two rows of cells along the midgut and in a single cell in each hemisegment of the CNS. Other sites include 1 to 2 cells in the dorsal part of each hemisegment from T to A6, cells along the pharyngeal muscle and along the hindgut. In late embryos, weaker expression is observed in somatic muscles.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
wing pouch

Comment: NOT presumptive wing veins, NOT dorsal/ventral compartment boundary

Additional Descriptive Data

bs expression was compared in the wing and haltere discs. Expression is present in the wing pouch but not in the haltere.

bs protein is expressed in the wing cell (intervein) region of the wing disc; expression is excluded from the presumptive wing veins and presumptive wing margin.

Expression of bs occurs in the intervein tissue of the developing pupal wing in a pattern complementary to that of ve.

Well defined staining of crossveins is observed by 23-26 hr APF.

bs protein is expressed ubiquitously in unfertilized eggs. The protein fades away during germ band extension and zygotic expression is not detected until germ band retraction. At stage 14, expression is observed just beneath the epidermis in a cluster of 6-9 cells in the most anterior portion of each hemisegment from T2 to A8. Expression is also observed in the head, in two rows of cells along the midgut and in a single cell in each hemisegment of the CNS. Other sites include 1 to 2 cells in the dorsal part of each hemisegment from T2 to A6, cells along the pharyngeal muscle and along the hindgut. In late embryos, weaker expression is observed in somatic muscles. Using double staining techniques, it was shown that all of the cells that express bs strongly are tightly associated with the tracheal system.

Marker for
Subcellular Localization
CV Term
Evidence
References
located_in nucleus
inferred from high throughput direct assay
inferred from direct assay
Expression Deduced from Reporters
Reporter: P{bs-GAL4.M}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{bs-GAL4.Term}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GAL4}bs1348
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GAL4-bs.23.26}
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\bs in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 57 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 15 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of bs
Transgenic constructs containing regulatory region of bs
Aberrations (Deficiencies and Duplications) ( 14 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
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
follicle cell & actin filament | somatic clone
wing & macrochaeta
wing vein L3 & sensillum campaniformium
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (6)
9 of 14
Yes
Yes
1  
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (5)
9 of 14
Yes
Yes
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
Mus musculus (laboratory mouse) (4)
9 of 14
Yes
Yes
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
Xenopus tropicalis (Western clawed frog) (3)
4 of 13
Yes
Yes
1 of 13
No
Yes
1 of 13
No
No
Danio rerio (Zebrafish) (8)
8 of 14
Yes
Yes
7 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (2)
8 of 14
Yes
Yes
2 of 14
No
No
Anopheles gambiae (African malaria mosquito) (3)
2 of 12
Yes
Yes
2 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (119)
3 of 13
Yes
No
3 of 13
Yes
No
3 of 13
Yes
No
3 of 13
Yes
No
3 of 13
Yes
No
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Yes
No
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Yes
No
2 of 13
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No
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Yes
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2 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
No
1 of 13
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No
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1 of 13
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Yes
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Yes
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Yes
1 of 13
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Yes
1 of 13
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Yes
1 of 13
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No
1 of 13
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No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
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Yes
1 of 13
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Yes
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Yes
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Yes
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Yes
1 of 13
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No
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No
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1 of 13
No
Yes
1 of 13
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No
1 of 13
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Yes
1 of 13
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Yes
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
Yes
1 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (4)
8 of 13
Yes
Yes
6 of 13
No
Yes
2 of 13
No
No
2 of 13
No
No
Schizosaccharomyces pombe (Fission yeast) (3)
5 of 12
Yes
Yes
2 of 12
No
No
1 of 12
No
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:bs. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (1)
2 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 1 )
    Allele
    Disease
    Interaction
    References
    Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    Functional Complementation Data
    Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
    Interactions
    Summary of Physical Interactions
    Summary of Genetic Interactions
    Interaction Browsers

    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
    suppressible
    suppressible
    suppressible
    External Data
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    FlyBase
    External Links
    External Data
    Linkouts
    Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2R
    Recombination map
    2-106
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    60C6-60C7
    Limits computationally determined from genome sequence between P{lacW}Nop60Bk05318 and P{PZ}bs03267&P{lacW}l(2)k10502k10502
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    60C6-60C8
    (determined by in situ hybridisation)
    60C-60C
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Location
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (38)
    Genomic Clones (24)
    cDNA Clones (20)
     

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

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

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

    cDNA Clones, End Sequenced (ESTs)
    RNAi and Array Information
    Linkouts
    DRSC - Results frm RNAi screens
    Antibody Information
    Laboratory Generated Antibodies
     
    Commercially Available Antibodies
     
    Cell Line Information
    Publicly Available Cell Lines
     
      Other Stable Cell Lines
       
        Other Comments

        bs encodes for two protein isoforms, one much longer than the other (449 vs. 355 amino acids). The long isoform is encoded by two transcripts (variants A and C), that use two different transcription start sites and differ only in the first exon. The other protein isoform is encoded by a shorter two-exon transcript (variant B), which shares its TSS with isoform C. The two long isoforms contain a 26 Kb intron, which holds the natural antisense transcript aslncRNA:bsAS.

        RNAi generated by PCR using primers directed to this gene causes a cell growth and viability phenotype when assayed in Kc167 and S2R+ cells.

        bs is involved in cell fate allocation and differentiation but does not participate in cell proliferation. The activity of bs is necessary to specify cells to adopt an intervein fate. It expression is repressed in larval stages in vein territories by genes of the Ras signalling pathway. The differentiation of vein and intervein depends on the outcome of a balance between bs and rho activities, achieved during pupal stages.

        bs regulates formation of the terminal branches of the tracheal system. bs may function like mammalian SRF in an inducible transcription complex. Activation of the complex by signals from target tissues induces expression of genes involved in cytoplasmic outgrowth.

        The bs gene product plays a dual role during wing differentiation. It acts in a dosage-dependent manner to suppress the formation of wing veins and is required cell-autonomously to promote the development of intervein cells. bs acts at a key step between regulatory genes that define the early positional values in the developing wing disc and the subsequent localized expression of intervein-specific structural genes.

        Required for the formation and outgrowth of the terminal branches of the tracheal system.

        The bs gene might play a role in the proper formation and maintenance of the trachea.

        Developmental and genetic analyses suggest that the bs product is required during metamorphosis for the initiation of intervein development and the concommitant inhibition of vein development.

        bs may affect dorsal-ventral flattening of the wing during prepupal morphogenesis.

        Relationship to Other Genes
        Source for database merge of
        Additional comments

        May be part of a pseudoallelic complex with Px.

        Nomenclature History
        Source for database identify of

        Source for identity of: bs CG3411

        Nomenclature comments
        Etymology
        Synonyms and Secondary IDs (38)
        Reported As
        Symbol Synonym
        Serf
        l(2)03267
        Name Synonyms
        Drosophila serum response
        Drosophila serum response factor
        blistered/DSRF
        blistered/Drosophila serum response factor
        Secondary FlyBase IDs
        • FBgn0000156
        • FBgn0000223
        • FBgn0010354
        • FBgn0010535
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 54 )
        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 Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
        GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
        RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
        UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
        UniProt/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
        AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
        BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
        DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
        EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
        FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
        FlyMine - An integrated database for Drosophila genomics
        KEGG Genes - Molecular building blocks of life in the genomic space.
        MARRVEL_MODEL - MARRVEL (model organism gene)
        Linkouts
        BioGRID - A database of protein and genetic interactions.
        Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
        DroID - A comprehensive database of gene and protein interactions.
        DRSC - Results frm RNAi screens
        Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
        FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
        FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
        Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
        Flygut - An atlas of the Drosophila adult midgut
        FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
        iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
        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.
        References (327)