FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\Z600
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General Information
Symbol
Dmel\Z600
Species
D. melanogaster
Name
Z600
Annotation Symbol
CG17962
Feature Type
protein_coding_gene
FlyBase ID
FBgn0004052
Gene Model Status
Current
Stock Availability
7 publicly available
Gene Summary
Z600 (Z600) encodes a mitotic inhibitor that binds to the hydrophobic patch of Cyclins. It contributes to the switch in cell cycle mode in blastoderm embryos, and delaying mitosis in the mesoderm anlage to avoid an interference of mitosis and morphogenesis. [Date last reviewed: 2019-03-21] (FlyBase Gene Snapshot)
All Summaries
Also Known As

frs, fruhstart, DMZ60MEX

Key Links
Genomic Location
Cytogenetic map
71D3-71D3
Sequence location
Recombination map
3-43
RefSeq locus
NT_037436 REGION:15509061..15509513
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 (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
enables cyclin binding
inferred from physical interaction with FLYBASE:CycA; FB:FBgn0000404
(Gawlinski et al., 2007)
inferred from physical interaction with FLYBASE:CycE; FB:FBgn0010382
(Gawlinski et al., 2007)
enables protein binding
inferred from physical interaction with FLYBASE:Nup50; FB:FBgn0033264
(Gawlinski et al., 2007)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
involved_in negative regulation of G2/M transition of mitotic cell cycle
inferred from mutant phenotype
(Gawlinski et al., 2007)
involved_in negative regulation of mitotic nuclear division
inferred from genetic interaction with FLYBASE:stg; FB:FBgn0003525
(Großhans and Wieschaus, 2000)
inferred from mutant phenotype
(Großhans et al., 2003)
involved_in ventral furrow formation
inferred from genetic interaction with FLYBASE:stg; FB:FBgn0003525
(Großhans and Wieschaus, 2000)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
involved_in gastrulation
non-traceable author statement
(Lee and Orr-Weaver, 2003, Packer, 2003)
involved_in negative regulation of mitotic nuclear division
non-traceable author statement
(Packer, 2003)
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
(Großhans et al., 2003)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
located_in cytoplasm
non-traceable author statement
(Packer, 2003)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
    -
    Summaries
    Gene Snapshot
    Z600 (Z600) encodes a mitotic inhibitor that binds to the hydrophobic patch of Cyclins. It contributes to the switch in cell cycle mode in blastoderm embryos, and delaying mitosis in the mesoderm anlage to avoid an interference of mitosis and morphogenesis. [Date last reviewed: 2019-03-21]
    Protein Function (UniProtKB)
    Cell cycle regulator that is involved in modulating and adjusting cell proliferation according to the requirements of the developmental program (PubMed:10850494, PubMed:12919679, PubMed:17431409). Interacts with mitotic Cdk1-cyclin complexes to inhibit mitotic entry at the G2/M transition (PubMed:17431409). Likely to function by binding to the hydrophobic patch of cyclins to interfere with the interaction between the complex and certain Cdk1 substrates (PubMed:17431409). At the mid-blastula transition, involved in the cell cycle arrest in G2 of cycle 14 by delaying mitosis and thus reducing cell proliferation allowing cell fate specification and morphogenesis to take place (PubMed:12919679). Acts downstream or in parallel to the checkpoint regulator grp which is also required for the cell cycle pause at cycle 14 (PubMed:12919679). During gastrulation, delays mitosis in the ventral region of the embryonic mesoderm thus allowing invagination to be completed before cell division takes place (PubMed:10850494, PubMed:12919679, PubMed:17431409).
    (UniProt, P22469)
    Phenotypic Description (Red Book; Lindsley and Zimm 1992)
    Z600
    Member of cluster of genes that includes Eip28/29 and Gdl. Believed to play role in processes of early development.
    Gene Model and Products
    Number of Transcripts
    1
    Number of Unique Polypeptides
    1

    Please see the JBrowse view of Dmel\Z600 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 P22469)

    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 5.44

    Gene model reviewed during 5.46

    Transcript Data
    Annotated Transcripts
    Name
    FlyBase ID
    RefSeq ID
    Length (nt)
    Assoc. CDS (aa)
    Z600-RA
    FBtr0075549
    NM_079359
    453
    90
    Additional Transcript Data and Comments
    Reported size (kB)

    0.6 (northern blot)

    (Schulz and Butler, 1989, Schulz and Miksch, 1989)
    Comments
    External Data
    Crossreferences
    Polypeptide Data
    Annotated Polypeptides
    Name
    FlyBase ID
    Predicted MW (kDa)
    Length (aa)
    Theoretical pI
    UniProt
    RefSeq ID
    GenBank
    Z600-PA
    FBpp0075304
    10.5
    90
    10.59
    P22469
    NP_524083
    AAF49634
    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)

    18 (kD observed); 10.5 (kD predicted)

    (Galewsky et al., 1990)
    Comments
    External Data
    Subunit Structure (UniProtKB)

    Component of the Frs-CycA-Cdk1 complex composed of Z600, CycA and Cdk1. Interacts preferentially with CycA (via C-terminus) but is also able to interact (via C-terminus) with CycE (via C-terminus).

    (UniProt, P22469)
    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\Z600 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).

    2.15

    Transcript Expression
    in situ
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    embryonic stage 4 -- 6
    organism | subset
    (Fisher et al., 2012)
    embryonic stage 7 -- 8
    amnioserosa anlage
    (Fisher et al., 2012)
    dorsal ectoderm
    (Fisher et al., 2012)
    embryonic cycle 14
    organism | ubiquitous
    (Sung et al., 2013)
    embryonic stage 9 -- 10
    amnioserosa primordium
    (Fisher et al., 2012)
    northern blot
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    embryonic stage 1 -- 9

    Comment: reference states 0-4 hr AEL

    (Schulz and Miksch, 1989)
    embryonic stage 4
    organism | dorsal

    Comment: late embryonic stage 4

    (Schulz and Miksch, 1989)
    organism | posterior

    Comment: late embryonic stage 4

    (Schulz and Miksch, 1989)
    embryonic cycle 13 -- embryonic cycle 16
    (Schulz and Miksch, 1989)
    embryonic stage 5 -- 12
    organism | dorsal
    (Schulz and Miksch, 1989)
    Additional Descriptive Data

    Z600 transcripts are detected starting at embryonic cycle 13. Transcripts initially accumulate uniformly, but become concentrated in the dorsal and posterior regions during cellularization, and persist in the dorsal regions of the embryo during gastrulation and germ band extension. No signal is detected after germ band extension.

    (Schulz and Miksch, 1989)
    Marker for
     
    Subcellular Localization
    CV Term
    Polypeptide Expression
    western blot
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    embryonic stage 1 -- 9

    Comment: reference states 0-4 hr AEL

    (Galewsky et al., 1990)
    embryonic cycle 11 -- embryonic cycle 13
    (Galewsky et al., 1990)
    Additional Descriptive Data

    The 18 kD Z600 protein is detected at low levels in 0-2 hr embryos and at higher levels in 2-4 hr embryos. No expression is detected in older embryos or in unfertilized eggs. Immunolocalization experiments reveal that expression is found only in nuclear cycle 11-13 embryos, associated with condensed chromatin. Protein is not detected at later stages, despite the presence of Z600 transcript.

    (Galewsky et al., 1990)
    Marker for
     
    Subcellular Localization
    CV Term
    Evidence
    References
    located_in cytoplasm
    inferred from direct assay
    (Großhans et al., 2003)
    Expression Deduced from Reporters
    Reporter: P{z600-lacZ}
    Stage
    Tissue/Position (including subcellular localization)
    Reference
    embryonic stage 1 -- 11

    Comment: reference states 0-6 hr AEL

    (Schulz and Miksch, 1989)
    embryonic stage 4
    organism | dorsal

    Comment: late embryonic stage 4

    (Schulz and Miksch, 1989)
    organism | posterior

    Comment: late embryonic stage 4

    (Schulz and Miksch, 1989)
    organism | ventral

    Comment: late embryonic stage 4

    (Schulz and Miksch, 1989)
    embryonic cycle 13 -- embryonic cycle 16
    (Schulz and Miksch, 1989)
    embryonic stage 5 -- 12
    organism | dorsal
    (Schulz and Miksch, 1989)
    organism | ventral
    (Schulz and Miksch, 1989)
    High-Throughput Expression Data
    Associated Tools

    JBrowse - Visual display of RNA-Seq signals

    View Dmel\Z600 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
    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 ( 12 )
    For All Alleles Carried on Transgenic Constructs Show
    Transgenic constructs containing/affecting coding region of Z600
    Transgenic constructs containing regulatory region of Z600
    Aberrations (Deficiencies and Duplications) ( 9 )
    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) (0)
    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) (0)
    Mus musculus (laboratory mouse) (0)
    Xenopus tropicalis (Western clawed frog) (0)
    Danio rerio (Zebrafish) (0)
    Caenorhabditis elegans (Nematode, roundworm) (0)
    Anopheles gambiae (African malaria mosquito) (0)
    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:Z600. 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
      Summary of Genetic Interactions
      Interaction Browsers
      View in FlyBase Interactions Browser   View in MIST tool (external link)
      Please look at the allele data for full details of the genetic interactions
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      Z600
      suppressible
      CycE
      (Gawlinski et al., 2007)
      Starting gene(s)
      Interaction type
      Interacting gene(s)
      Reference
      Cdk1
      suppressible
      Z600
      (Gawlinski et al., 2007)
      External Data
      Subunit Structure (UniProtKB)
      Component of the Frs-CycA-Cdk1 complex composed of Z600, CycA and Cdk1. Interacts preferentially with CycA (via C-terminus) but is also able to interact (via C-terminus) with CycE (via C-terminus).
      (UniProt, P22469 )
      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
      3-43
      Cytogenetic map
      71D3-71D3
      Sequence location
      FlyBase Computed Cytological Location
      Cytogenetic map
      Evidence for location
      71D3-71D3
      Limits computationally determined from genome sequence between P{lacW}cp309s2172&P{lacW}l(3)j2A2j2A2 and P{EP}Tfb2EP572
      Experimentally Determined Cytological Location
      Cytogenetic map
      Notes
      References
      71C-71C
      (Gibson et al., 1999)
      Experimentally Determined Recombination Data
      Location

      3-43

      (FlyBase, 2016)

      3-37.2

      (Comeron et al., 2012)
      Left of (cM)
      Right of (cM)
      Notes
      Stocks and Reagents
      Stocks (7)
      Genomic Clones (16)
       

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

      cDNA Clones (22)
       

      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

          (Galewsky et al., 1990)
          Commercially Available Antibodies
           
          Cell Line Information
          Publicly Available Cell Lines
           
            Other Stable Cell Lines
             
              Other Comments

              Z600 controls the cell cycle pause after precisely thirteen mitoses in embryos.

              (Großhans et al., 2003)

              In embryos lacking Z600 function, cells in the ventral domain enter mitosis too early, causing a defective ventral furrow.

              (Großhans et al., 2003)

              Z600 is required for the ventral inhibition of mitosis that acts during gastrulation to counteract stg specifically in the cells that invaginate to form the mesoderm.

              (Großhans and Wieschaus, 2000)

              In embryos lacking Z600 function, cells in the ventral domain are the first to enter mitosis, such that their pattern of stg RNA expression and the mitotic pattern match each other.

              (Großhans and Wieschaus, 2000)

              Variation of a microsatellite within the Z600 locus has been studied in North American populations of D.melanogaster.

              (Goldstein and Clark, 1995)

              The pattern of Z600 protein expression during early embryogenesis has been determined.

              (Galewsky et al., 1990)

              The spatial and temporal expression of Z600 RNA has been studied.

              (Schulz and Miksch, 1989)

              Identification: Part of an analysis of Eip71CD gene expression.

              (Schulz et al., 1989)

              Z600 has been identified and characterised.

              (Schulz et al., 1989)

              Member of cluster of genes that includes Eip28/29 and gdl. Believed to play role in processes of early development.

              Relationship to Other Genes
              Source for database merge of

              Source for merge of: Z600 frs

              (Großhans et al., 2003)
              Additional comments
              Nomenclature History
              Source for database identify of
              Nomenclature comments
              Etymology
              Synonyms and Secondary IDs (10)
              Reported As
              Symbol Synonym
              3.2
              (Gibson et al., 1999)
              CG17962
              (Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Lu et al., 2009, Brandt, 2006)
              DMZ60MEX
              (Fernando Vazquez et al., 2000, Schug et al., 1997)
              Frs
              (Großhans et al., 2007)
              Z600
              (Örkenby et al., 2023, Transgenic RNAi Project members, 2017-, Chen et al., 2013, Saunders et al., 2013, Liang et al., 2008, Reiter et al., 2007, Sandmann et al., 2007)
              frs
              (Syed et al., 2021, Sung et al., 2013, Nien et al., 2011, Papoulas et al., 2010, Insco et al., 2009, Insco et al., 2009, Lu et al., 2009, Liang et al., 2008, Gawlinski et al., 2007, Gawlinski et al., 2007, Sung and Großhans, 2007, Brandt, 2006, Pilot et al., 2006, Martin and Parkhurst, 2004, Lee and Orr-Weaver, 2003, Packer, 2003, Ip and Gridley, 2002, Greaves, 2000, Großhans and Wieschaus, 2000, Großhans and Wieschaus, 2000, Johnston, 2000)
              z600
              (Heimiller et al., 2014, Melcher and Pankratz, 2005, Großhans et al., 2003, Cherbas and Cherbas, 1997, Goldstein and Clark, 1995, Andres and Cherbas, 1994, Maroni, 1994, Cherbas, 1993, Cherbas et al., 1991, Cherbas, 1990.10.29, Galewsky et al., 1990, Schulz et al., 1990, Schulz et al., 1990, Schulz and Butler, 1989, Schulz and Miksch, 1989, Schulz et al., 1989)
              Name Synonyms
              Fruhstart
              (Gawlinski et al., 2007)
              Z600
              fruhstart
              (Liu and Grosshans, 2017, Yokoyama and Nakamura, 2011, Papoulas et al., 2010, Insco et al., 2009, Lu et al., 2009, Großhans et al., 2007, Pilot et al., 2006, Großhans and Wieschaus, 2000)
              frustart
              (Johnston, 2000)
              Secondary FlyBase IDs
              • FBgn0026799
              • FBgn0041755
              Datasets (0)
              Study focus (0)
              Experimental Role
              Project
              Project Type
              Title
              Study result (0)
              Result
              Result Type
              Title
              External Crossreferences and Linkouts ( 25 )
              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
              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
              Flygut - An atlas of the Drosophila adult midgut
              MIST (protein-protein) - An integrated Molecular Interaction Database
              References (76)