FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\Mis12
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General Information
Symbol
Dmel\Mis12
Species
D. melanogaster
Name
Mis12
Annotation Symbol
CG18156
Feature Type
protein_coding_gene
FlyBase ID
FBgn0035725
Gene Model Status
Current
Stock Availability
13 publicly available
Gene Summary
Mis12 (Mis12) encodes a protein involved in chromosome movement and metaphase plate congression during mitosis. [Date last reviewed: 2019-09-26] (FlyBase Gene Snapshot)
All Summaries
Also Known As

dmMis12

Key Links
Genomic Location
Cytogenetic map
65D5-65D5
Sequence location
Recombination map
3-18
RefSeq locus
NT_037436 REGION:6968035..6969048
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (7 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables kinetochore adaptor activity
inferred from direct assay
(Richter et al., 2016)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
involved_in chromosome segregation
inferred from mutant phenotype
(Przewloka et al., 2007)
involved_in mitotic chromosome movement towards spindle pole
inferred from mutant phenotype
(Venkei et al., 2011)
involved_in mitotic metaphase chromosome alignment
inferred from mutant phenotype
(Przewloka et al., 2007)
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
located_in chromosome, centromeric region
inferred from direct assay
(Venkei et al., 2011)
located_in kinetochore
inferred from direct assay
(Venkei et al., 2011, Goshima et al., 2007, Przewloka et al., 2007, Schittenhelm et al., 2007)
part_of MIS12/MIND type complex
inferred from direct assay
(Richter et al., 2016, Schittenhelm et al., 2007)
Terms Based on Predictions or Assertions (0 terms)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
    -
    Summaries
    Gene Snapshot
    Mis12 (Mis12) encodes a protein involved in chromosome movement and metaphase plate congression during mitosis. [Date last reviewed: 2019-09-26]
    Gene Group (FlyBase)
    KINETOCHORE MIS12 COMPLEX - NF1A VARIANT -
    The MIS12 complex is an outer kinetochore complex that bridges the inner kinetochore component CENP-C and the outer kinetochore Ndc80 complex. In Drosophila, there are two complex variants, one containing Nnf1a and one containing Nnf1b. (Adapted from FBrf0231098.)
    KINETOCHORE MIS12 COMPLEX - NF1B VARIANT -
    The MIS12 complex is an outer kinetochore complex that bridges the inner kinetochore component CENP-C and the outer kinetochore Ndc80 complex. In Drosophila, there are two complex variants, one containing Nnf1a and one containing Nnf1b. (Adapted from FBrf0231098.)
    Gene Model and Products
    Number of Transcripts
    1
    Number of Unique Polypeptides
    1

    Please see the JBrowse view of Dmel\Mis12 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 Q9VS01)

    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 5.46

    Transcript Data
    Annotated Transcripts
    Name
    FlyBase ID
    RefSeq ID
    Length (nt)
    Assoc. CDS (aa)
    Mis12-RA
    FBtr0076936
    NM_139809
    839
    181
    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
    Mis12-PA
    FBpp0076645
    21.1
    181
    5.98
    Q9VS01
    NP_648066
    AAF50629
    Polypeptides with Identical Sequences

    There is only one protein coding transcript and one polypeptide associated with this gene

    Additional Polypeptide Data and Comments
    Reported size (kDa)
    Comments
    External Data
    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\Mis12 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).

    2.83

    Transcript Expression
    in situ
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    embryonic stage 1 -- 3
    organism

    Comment: maternally deposited

    (Fisher et al., 2012)
    Additional Descriptive Data
    Marker for
     
    Subcellular Localization
    CV Term
    Polypeptide Expression
    epitope tag
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    embryonic stage 4 -- 5
    cell

    Comment: Localization observed prometaphase until late anaphase of embryonic cycle 14

    (Schittenhelm et al., 2007)
    cell

    Comment: Localization observed in mitotic cells; assayed in cultured S2R+ cells

    (Schittenhelm et al., 2007)
    Additional Descriptive Data

    Subcellular localization to kinetochores of embryonic mitotic cells assayed using the transgenic allele Mis12T:AvicGFP-EGFP, and of cultured S2R+ cells using the transgenic allele Mis12Scer\UAS.T:AvicGFP-EGFP.

    (Schittenhelm et al., 2007)
    Marker for
     
    Subcellular Localization
    CV Term
    Evidence
    References
    located_in chromosome, centromeric region
    inferred from direct assay
    (Venkei et al., 2011)
    located_in kinetochore
    inferred from direct assay
    (Venkei et al., 2011, Goshima et al., 2007, Przewloka et al., 2007, Schittenhelm et al., 2007)
    part_of MIS12/MIND type complex
    inferred from direct assay
    (Richter et al., 2016, Schittenhelm et al., 2007)
    Expression Deduced from Reporters
    Reporter: P{gMis12-EGFP}
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    embryonic stage 4 -- 5
    cell

    Comment: Localization observed prometaphase until late anaphase of embryonic cycle 14

    (Schittenhelm et al., 2007)
    High-Throughput Expression Data
    Associated Tools

    JBrowse - Visual display of RNA-Seq signals

    View Dmel\Mis12 in JBrowse
    RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
    Bulk Downloads
    RNA-Seq RPKM values for all genes
    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
    FlyBase Wiki
    Image Based Resources
    Alleles, Insertions, Transgenic Constructs, and Aberrations
    Classical and Insertion Alleles ( 2 )
    For All Classical and Insertion Alleles Show
     
    Other relevant insertions
    Transgenic Constructs ( 11 )
    For All Alleles Carried on Transgenic Constructs Show
    Transgenic constructs containing/affecting coding region of Mis12
    Transgenic constructs containing regulatory region of Mis12
    Aberrations (Deficiencies and Duplications) ( 1 )
    Inferred from experimentation ( 1 )
    Gene partially disrupted in
    Df(3L)vvl7247-Mis12
    (Sotillos et al., 2010)
    Inferred from location ( 5 )
    Variants
    Variant Molecular Consequences
    Alleles Representing Disease-Implicated Variants
    Phenotypes
    Orthologs
    Downloads
    Download All DIOPT Orthologs
    Human Orthologs (via DIOPT v9.1)
    Species\Gene Symbol
    Score
    Best Score
    Best Reverse Score
    Source
    Compara
    Domainoid
    eggNOG
    Hieranoid
    Homologene
    Inparanoid
    OMA
    OrthoDB
    OrthoFinder
    OrthoInspector
    orthoMCL
    Panther
    Phylome
    SonicParanoid
    Alignment
    Complementation?
    Transgene?
    Homo sapiens (Human) (1)
    Hsap\MIS12
    1 of 14
    Yes
    Yes
    Model Organism Orthologs (via DIOPT v9.1)
    Species\Gene Symbol
    Score
    Best Score
    Best Reverse Score
    Source
    Compara
    Domainoid
    eggNOG
    Hieranoid
    Homologene
    Inparanoid
    OMA
    OrthoDB
    OrthoFinder
    OrthoInspector
    orthoMCL
    Panther
    Phylome
    SonicParanoid
    Alignment
    Complementation?
    Transgene?
    Rattus norvegicus (Norway rat) (1)
    Rnor\Mis12
    1 of 14
    Yes
    Yes
    Mus musculus (laboratory mouse) (1)
    Mmus\Mis12
    1 of 14
    Yes
    Yes
    Xenopus tropicalis (Western clawed frog) (0)
    Danio rerio (Zebrafish) (1)
    Drer\mis12
    1 of 14
    Yes
    Yes
    Caenorhabditis elegans (Nematode, roundworm) (1)
    Cele\mis-12
    1 of 14
    Yes
    Yes
    Anopheles gambiae (African malaria mosquito) (0)
    Arabidopsis thaliana (thale-cress) (0)
    Saccharomyces cerevisiae (Brewer's yeast) (0)
    Schizosaccharomyces pombe (Fission yeast) (1)
    Spom\mis12
    1 of 12
    Yes
    Yes
    Escherichia coli (enterobacterium) (0)
    Other Organism Orthologs (via OrthoDB)
    Data provided directly from OrthoDB:Mis12. Refer to their site for version information.
    Paralogs
    Downloads
    Download All DIOPT Paralogs
    Paralogs (via DIOPT v9.1)
    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 ( 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
      Interaction Browsers
      View in FlyBase Interactions Browser   View in MIST tool (external link)
      Please see the Physical Interaction reports below for full details
      protein-protein
      Physical Interaction
      Assay
      References
      Mis12 - Atac1
      two hybrid
      (Hu et al., 2017.6.13)
      Mis12 - CG3339
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Przewloka et al., 2007)
      Mis12 - Cenp-C
      mass spectrometry study of hydrogen/deuterium exchange, molecular sieving, cosedimentation, molecular weight estimation by staining, anti tag coimmunoprecipitation, peptide massfingerprinting, anti bait coimmunoprecipitation, pull down, protein cross-linking with a bifunctional reagent
      (Liu et al., 2016, Richter et al., 2016, Przewloka et al., 2011)
      Mis12 - Dhc16F
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Przewloka et al., 2007)
      Mis12 - Dhc62B
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Przewloka et al., 2007)
      Mis12 - Dhc64C
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Przewloka et al., 2007)
      Mis12 - Kmn1
      anti bait coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based, two hybrid, anti tag coimmunoprecipitation, pull down, molecular sieving, mass spectrometry study of hydrogen/deuterium exchange, cosedimentation, molecular weight estimation by staining, protein cross-linking with a bifunctional reagent
      (Liu et al., 2016, Richter et al., 2016, Guruharsha et al., 2011, Schittenhelm et al., 2009, Przewloka et al., 2007)
      Mis12 - Ndc80
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Przewloka et al., 2007)
      Mis12 - Nnf1a
      anti tag coimmunoprecipitation, peptide massfingerprinting, molecular sieving, cosedimentation, molecular weight estimation by staining, pull down, anti bait coimmunoprecipitation, two hybrid, experimental knowledge based, protein cross-linking with a bifunctional reagent, anti tag western blot, mass spectrometry study of hydrogen/deuterium exchange
      (Liu et al., 2016, Richter et al., 2016, Guruharsha et al., 2011, Schittenhelm et al., 2009, Przewloka et al., 2007)
      Mis12 - Nnf1b
      pull down, anti tag western blot, molecular sieving, cosedimentation, molecular weight estimation by staining, two hybrid, protein cross-linking with a bifunctional reagent, peptide massfingerprinting, experimental knowledge based, anti tag coimmunoprecipitation
      (Hu et al., 2017.6.13, Liu et al., 2016, Guruharsha et al., 2011, Przewloka et al., 2007)
      Mis12 - Nuf2
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Przewloka et al., 2007, Schittenhelm et al., 2007)
      Mis12 - Spc105R
      molecular sieving, cosedimentation, molecular weight estimation by staining, anti tag coimmunoprecipitation, peptide massfingerprinting
      (Liu et al., 2016, Przewloka et al., 2007)
      Mis12 - Spc25
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Przewloka et al., 2007, Schittenhelm et al., 2007)
      Mis12 - btv
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Przewloka et al., 2007)
      Mis12 - cana
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Przewloka et al., 2007)
      Mis12 - rod
      anti tag coimmunoprecipitation, peptide massfingerprinting
      (Przewloka et al., 2007)
      Summary of Genetic Interactions
      Interaction Browsers
      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 (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)
      3L
      Recombination map
      3-18
      Cytogenetic map
      65D5-65D5
      Sequence location
      FlyBase Computed Cytological Location
      Cytogenetic map
      Evidence for location
      65D5-65D5
      Limits computationally determined from genome sequence between P{lacW}l(3)L4060L4060 and P{PZ}corn04202
      Experimentally Determined Cytological Location
      Cytogenetic map
      Notes
      References
      Experimentally Determined Recombination Data
      Location

      3-18

      (FlyBase, 2016)

      3-15.8

      (Comeron et al., 2012)
      Left of (cM)
      Right of (cM)
      Notes
      Stocks and Reagents
      Stocks (13)
      Genomic Clones (11)
       

      Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

      cDNA Clones (4)
       

      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
        Other clones
        RNAi and Array Information
        Linkouts
        DRSC - Results frm RNAi screens
        Antibody Information
        Laboratory Generated Antibodies
         

        polyclonal

        (Venkei et al., 2011)
        Commercially Available Antibodies
         
        Cell Line Information
        Publicly Available Cell Lines
         
          Other Stable Cell Lines
           

          • New stable cell line derived from S2-unspecified : A stable cell line containing a fusion of Mis12 and human Katanin p60 to specifically localize Katanin p60 to the kinetochore was produced.

            (Advani et al., 2018)
          Other Comments

          RNAi screen using dsRNA made from templates generated with primers directed against this gene results in chromosome misalignment on the metaphase spindle and a spindle that is aberrantly long when assayed in S2 cells. This phenotype can be observed when the screen is performed with or without Cdc27 dsRNA.

          (Goshima et al., 2007)
          Relationship to Other Genes
          Source for database merge of

          Source for merge of: CG18156 BcDNA:RE19545

          (FlyBase, 1996-)
          Additional comments

          FlyBase curator comment: Correspondence with author Lehner revealed that 'Mis12' = 'CG18156'.

          (Lehner et al., 2007)

          Source for merge of CG18156 BcDNA:RE19545 was a shared cDNA ( date:020730 ).

          (FlyBase, 1996-)
          Nomenclature History
          Source for database identify of

          Source for identity of: Mis12 CG18156

          (Goshima et al., 2007, Przewloka et al., 2007, Schittenhelm et al., 2007)
          Nomenclature comments
          Etymology
          Synonyms and Secondary IDs (8)
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 28 )
          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
          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
          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
          MIST (protein-protein) - An integrated Molecular Interaction Database
          References (70)