FB2025_01 , released February 20, 2025
Gene: Dmel\mid
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General Information
Symbol
Dmel\mid
Species
D. melanogaster
Name
midline
Annotation Symbol
CG6634
Feature Type
FlyBase ID
FBgn0261963
Gene Model Status
Stock Availability
Gene Summary
T-box transcription factors that participate in cardiac fate specificity - regulates wing development by repressing wingless and hedgehog (Interactive Fly, mid )
Contribute a Gene Snapshot for this gene.
Also Known As

nmr2, extra, neuromancer2, los, lost in space

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
2-17
RefSeq locus
NT_033779 REGION:5461639..5467610
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (32 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
Biological Process (24 terms)
Terms Based on Experimental Evidence (21 terms)
CV Term
Evidence
References
involved_in axon guidance
inferred from mutant phenotype
involved_in axonogenesis
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:HGTX; FB:FBgn0040318
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
involved_in gonad development
inferred from mutant phenotype
involved_in heart development
inferred from mutant phenotype
involved_in heart process
inferred from direct assay
inferred from genetic interaction with FLYBASE:H15; FB:FBgn0016660
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:HGTX; FB:FBgn0040318
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:H15; FB:FBgn0016660
inferred from mutant phenotype
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000137774
involved_in heart development
inferred from biological aspect of ancestor with PANTHER:PTN002601330
Cellular Component (2 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in chromatin
inferred from biological aspect of ancestor with PANTHER:PTN004251338
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN004251338
located_in nucleus
inferred from electronic annotation with InterPro:IPR018186, InterPro:IPR036960
Gene Group (FlyBase)
Protein Family (UniProt)
-
Summaries
Gene Group (FlyBase)
T-BOX TRANSCRIPTION FACTORS -
The T-box transcription factors are sequence-specific DNA binding proteins that regulate transcription. They are characterized by a conserved DNA-binding immunoglobulin-fold domain of 180-200 amino acids, known as T-box, named after its founding member, the mouse T protein. (Adapted from FBrf0218841).
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
mid: midline
Homozygous embryonic lethal; denticle bands defective in ventral midline.
Summary (Interactive Fly)

T-box transcription factors that participate in cardiac fate specificity - regulates wing development by repressing wingless and hedgehog

Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\mid 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 Q9VMR2)

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)
FBtr0079093
2920
580
FBtr0310358
2139
580
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
FBpp0078726
62.9
580
8.31
FBpp0302039
62.9
580
8.31
Polypeptides with Identical Sequences

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

580 aa isoforms: mid-PA, mid-PB
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
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\mid 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).

-0.66

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
RT-PCR
Stage
Tissue/Position (including subcellular localization)
Reference
single cell RNA-seq
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

mid transcript is expressed in the developing dorsal vessel from embryonic stage 11.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

In wild-type third instar larvae, mid is ubiquitously expressed in the nucleus in both peripodial epithelium and the disc proper.

mid staining is found only to the posterior of the dorsal appendage forming follicle cells, where br expression is present but much weaker than in the roof cells. It is also not expressed in the br-negative cells on the dorsal midline, between the two patches of roof cells.

mid protein is expressed in a large number of neurons in the ventral midline. It was localized to specific midline neuroblasts at embryonic stages 10 and 11. At stage 13, it was found to be expressed in a row 5 GMC, dubbed an "M-neuron", that appears to change into an extra RP2. It's relation to the NB4-2->GMC-1->RP2/sib lineage is described.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\mid 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
Images
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 20 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 18 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of mid
Transgenic constructs containing regulatory region of mid
Aberrations (Deficiencies and Duplications) ( 27 )
Inferred from experimentation ( 27 )
Gene duplicated in
Inferred from location ( 4 )
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
Other Phenotypes
Allele
Phenotype manifest in
Allele
chordotonal organ & axon | lateral
denticle row 1 & abdominal 1 ventral denticle belt
denticle row 1 & abdominal 2 ventral denticle belt
denticle row 1 & abdominal 3 ventral denticle belt
denticle row 1 & abdominal 4 ventral denticle belt
denticle row 1 & abdominal 5 ventral denticle belt
denticle row 1 & abdominal 6 ventral denticle belt
denticle row 1 & abdominal 7 ventral denticle belt
denticle row 1 & abdominal 8 ventral denticle belt
denticle row 2 & abdominal 1 ventral denticle belt
denticle row 2 & abdominal 2 ventral denticle belt
denticle row 2 & abdominal 3 ventral denticle belt
denticle row 2 & abdominal 4 ventral denticle belt
denticle row 2 & abdominal 5 ventral denticle belt
denticle row 2 & abdominal 6 ventral denticle belt
denticle row 2 & abdominal 7 ventral denticle belt
denticle row 2 & abdominal 8 ventral denticle belt
denticle row 3 & abdominal 1 ventral denticle belt
denticle row 3 & abdominal 2 ventral denticle belt
denticle row 3 & abdominal 3 ventral denticle belt
denticle row 3 & abdominal 4 ventral denticle belt
denticle row 3 & abdominal 5 ventral denticle belt
denticle row 3 & abdominal 6 ventral denticle belt
denticle row 3 & abdominal 7 ventral denticle belt
denticle row 3 & abdominal 8 ventral denticle belt
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (17)
11 of 14
Yes
Yes
4 of 14
No
Yes
4 of 14
No
No
4 of 14
No
No
3 of 14
No
Yes
3 of 14
No
Yes
3 of 14
No
No
3 of 14
No
No
4  
3 of 14
No
No
5  
3 of 14
No
No
3 of 14
No
No
1  
3 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (17)
12 of 14
Yes
Yes
4 of 14
No
Yes
4 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
3 of 14
No
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
2 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Mus musculus (laboratory mouse) (17)
11 of 14
Yes
Yes
4 of 14
No
Yes
4 of 14
No
No
1  
3 of 14
No
Yes
3 of 14
No
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
3 of 14
No
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
2 of 14
No
No
2 of 14
No
No
Xenopus tropicalis (Western clawed frog) (23)
8 of 13
Yes
Yes
2 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 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
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Danio rerio (Zebrafish) (25)
10 of 14
Yes
Yes
4 of 14
No
No
4 of 14
No
No
4 of 14
No
No
3 of 14
No
Yes
3 of 14
No
Yes
3 of 14
No
Yes
3 of 14
No
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
3 of 14
No
Yes
3 of 14
No
No
3 of 14
No
No
3 of 14
No
Yes
3 of 14
No
No
2 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (22)
9 of 14
Yes
Yes
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
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
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
2 of 14
No
No
2 of 14
No
No
2 of 14
No
No
2 of 14
No
Yes
2 of 14
No
No
2 of 14
No
No
Anopheles gambiae (African malaria mosquito) (12)
11 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:mid. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (8)
10 of 13
5 of 13
4 of 13
4 of 13
4 of 13
4 of 13
4 of 13
3 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Allele
Disease
Evidence
References
Potential Models Based on Orthology ( 3 )
Modifiers Based on Experimental Evidence ( 0 )
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
esyN Network Diagram
Show neighbor-neighbor interactions:
Show/hide secondary interactors 
(data from AllianceMine provided by esyN)
Select Layout:
Legend:
Protein
RNA
Selected Interactor(s)
Other Interaction Browsers

Please see the Physical Interaction reports below for full details
protein-protein
Physical Interaction
Assay
References
Summary of Genetic Interactions
esyN Network Diagram
Show/hide secondary interactors 
(data from AllianceMine provided by esyN)
esyN Network Key:
Suppression
Enhancement
Other 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
External Data
Linkouts
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
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
2L
Recombination map
2-17
Cytogenetic map
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
25E2-25E2
Limits computationally determined from genome sequence between P{lacW}vrik05901&P{PZ}cype03771 and P{lacW}Hel25Ek11511
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
Experimentally Determined Recombination Data
Left of (cM)
Notes
Stocks and Reagents
Stocks (31)
Genomic Clones (19)
cDNA Clones (112)
 

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)
    BDGP DGC clones
    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

        mid is required for the formation and specification of neuroblasts in the anterior-most part of each embryonic segment.

        mid and H15 are negative regulators of wg expression and act to break the symmetry of hh signaling during segmentation of the embryo.

        Loss of both mid and H15 produces defects that mimic a wg gain-of-function phenotype.

        Mutants have an extra RP2 or RP2Sib neuron in the endogenous position as well as an ectopic RP2 neuron at the periphery of the nerve cord.

        Mutations in 12 complementation groups differentially affect lateral chordotonal axon growth, fasciculation or ventral orientation. Mutations in robo, spen, sli and los cause lch axon defasciculation. Mutations in sli and los also cause some lch axon bundles to grow dorsally along a trajectory 180o from normal.

        mid mutations inhibit the formation of axon tracts between segments of the embryo, sensory axon pathfinding in the PNS and certain tracheal and glial cell migrations.

        Mutations in mid generate malformations of the longitudinal tracts.

        The role of mid in the regulation of run mRNA expression in the early embryo has been investigated.

        Relationship to Other Genes
        Source for database merge of

        Source for merge of: nmr2 mid

        Source for merge of: mid nmr2

        Source for merge of: mid los extra

        Additional comments
        Nomenclature History
        Source for database identify of

        Source for identity of: nmr2 CG6634

        Nomenclature comments
        Etymology
        Synonyms and Secondary IDs (19)
        Reported As
        Symbol Synonym
        extra
        Secondary FlyBase IDs
        • FBgn0005511
        • FBgn0001934
        • FBgn0002746
        • FBgn0031706
        • FBgn0016996
        • FBgn0065989
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 37 )
        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/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
        KEGG Genes - Molecular building blocks of life in the genomic space.
        MARRVEL_MODEL - MARRVEL (model organism gene)
        Linkouts
        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
        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
        References (140)