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General Information
Symbol
Dmel\zen
Species
D. melanogaster
Name
zerknullt
Annotation Symbol
CG1046
Feature Type
FlyBase ID
FBgn0004053
Gene Model Status
Stock Availability
Gene Snapshot
In progress.Contributions welcome.
Also Known As
z1, zerknult
Key Links
Genomic Location
Cytogenetic map
Sequence location
3R:6,752,864..6,754,194 [-]
Recombination map
3-48
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Summaries
Gene Group (FlyBase)
HOX-LIKE HOMEOBOX TRANSCRIPTION FACTORS -
HOX-like (HOXL) homeobox transcription factors are sequence-specific DNA binding proteins that regulate transcription. They encompass transcription factors encoded by the Hox genes of the Antennapedia and the Bithorax gene complexes and genes closely related in sequence. HOXL transcription factors are major regulators of animal development. (Adapted from FBrf0232555).
Protein Function (UniProtKB)
Required for the differentiation of the dorsal-ventral pattern, and does not appear to be involved in the process of segmentation.
(UniProt, P09089)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
zen: zerknullt
Null mutations result in embryonic lethality and the loss of several dorsally derived embryonic structures, including the amnioserosa, optic lobe, and dorsal ridge. These animals also fail to fully extend their germ bands and go through the process of head involution. The name for the locus derives from the characteristic "wrinkled" appearance of the germ band seen in the SEM at the time of normal germ-band retraction. Hypomorphic mutations result in the absence of dorsal structures but do undergo normal gastrulation movements. A temperature-sensitive allele has been used to define the time of zen+ action between 2 and 4 hours of embryogenesis, just prior to and overlapping the earliest observable morphogenic defects. X-ray induced somatic clones have further shown that zen+ function is unnecessary for postembryonic development. The RNA product of zen is first detected at about 2 hours of development during the eleventh to twelfth cell cycle of the syncytial blastoderm. At this early stage the RNA is found on the dorsal surface of the embryo extending around the anterior and posterior poles. As cellularization proceeds and the early events of gastrulation begin, the RNA becomes restricted to a mid-dorsal stripe of cells. These cells have been fate mapped and give rise to the amnioserosa and the lobes in the dorsal posterior of the embryonic head, i.e., the structures absent in zen- animals. The time of appearance of zen RNA also correlates nicely with the temperature-sensitive-period data obtained using the conditional allele. Antisera to the zen protein product has been used to follow its accumulation pattern, and this analysis agrees with and expands the in situ results. The protein is located in the nuclei of cells expressing the gene and at cellular blastoderm is found in a mid-dorsal stripe seven cells wide and seventy cells in length. During gastrulation these cells eventually give rise to the amnioserosa, the optic lobe, and dorsal ridge; these structures continue to show zen protein accumulation until the end of germ-band extension at about 4 to 6 hours of development. This end point also correlates well with the end of the temperature-sensitive period of the conditional allele. The spatial pattern of zen expression has been shown to be dependent on the products of several of the maternally expressed genes which specify the anterior-posterior and dorsal-ventral polarity of the embryo, and zen would appear to lie near the end of the axis-determining pathway.
Summary (Interactive Fly)
transcription factor - homeodomain - Antp class - HoxA3, HoxB3 and HoxD3 homolog - DV polarity - The absence of the amnioserosa in mutants might cause the germ band to be twisted and thrown into folds during an abortive elongation process
Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\zen or the JBrowse view of Dmel\zen for information on other features

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

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Comments on Gene Model
Gene model reviewed during 5.49
Gene model reviewed during 5.50
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0081670
1267
353
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
RefSeq ID
GenBank
FBpp0081170
39.3
353
9.58
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)
353 (aa); 39 (kD predicted)
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\zen using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (8 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN002518688
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN002518688
(assigned by GO_Central )
Biological Process (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN002518688
(assigned by GO_Central )
Cellular Component (1 term)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN002518688
(assigned by GO_Central )
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
dorsal ectoderm anlage

Comment: anlage in statu nascendi

dorsal head epidermis anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

posterior ectoderm anlage

Comment: anlage in statu nascendi

antennal anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

visual anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

antennal anlage

Comment: reported as procephalic ectoderm anlage

central brain anlage

Comment: reported as procephalic ectoderm anlage

dorsal head epidermis anlage

Comment: reported as procephalic ectoderm anlage

visual anlage

Comment: reported as procephalic ectoderm anlage

organism | anterior | dorsal | posterior

Comment: anterior and posterior poles and dorsal surface of embryo

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
radioisotope in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism | dorsal

Comment: includes anterior and posterior pole

Additional Descriptive Data
Reporter constructs with deletions of 1.6 kb of zen upstream region driving lacZ expression demonstrate that a ventral repression element lies at about -1.2 to -1.4.
Northern analysis of RNA from 0-24 hour embryos indicate that zen transcript is expressed in early embryos, and that expression peaks at 2-3 hours of embryogenesis. Using radioactive in situ hybridization, zen expression is detected in the dorsal surface and the anterior and posterior poles of embryonic cycle 14 embryos. By gastrulation, zen expression is restricted to the dorsal surface. During germ band elongation, zen transcript is detected in the amnioserosa and a region of the dorsal ectoderm corresponding to the presumptive optic lobe, and in a subset of pole cells within the posterior midgut invagination.
The zen-RA transcripts are detected along the dorsal surface in embryos following the formation of the syncytial blastoderm. Following gastrulation, the expression of zen-RA is limited to a 5-6 cell wide stripe along the dorsal surface, and is not detected in the posterior 20% of the embryo.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Just prior to cellularization, zen protein is detected in the dorsal ectoderm. In later stages, during gastrulation, zen protein expression is resticted to a narrow dorsal stripe.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
Reporter: P{0.2zen-lacZ}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{0.7}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{1.0zen-lacZ}
Stage
Tissue/Position (including subcellular localization)
Reference
organism | dorsal | ventral

Comment: excludes anterior and posterior pole

Reporter: P{1.3}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{1.4}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{1.6zen-lacZ}
Stage
Tissue/Position (including subcellular localization)
Reference
organism | dorsal

Comment: includes anterior and posterior pole

organism | dorsal

Comment: also small amount of ventral expression

Reporter: P{ΔAcc}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{ΔNru}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{ΔRsa}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{ΔXmn}
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\zen in GBrowse 2
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 4-6
  • Stages(s) 7-8
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 10 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 20 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of zen
Transgenic constructs containing regulatory region of zen
Deletions and Duplications ( 31 )
Phenotypes
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (33)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
3 of 15
Yes
Yes
3 of 15
Yes
Yes
3 of 15
Yes
Yes
2 of 15
No
No
2 of 15
No
No
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
 
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (33)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
3 of 15
Yes
Yes
3 of 15
Yes
Yes
 
3 of 15
Yes
Yes
2 of 15
No
No
2 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Rattus norvegicus (Norway rat) (36)
3 of 13
Yes
Yes
3 of 13
Yes
Yes
3 of 13
Yes
Yes
2 of 13
No
No
2 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
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
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
Xenopus tropicalis (Western clawed frog) (26)
3 of 12
Yes
Yes
3 of 12
Yes
Yes
3 of 12
Yes
Yes
3 of 12
Yes
Yes
2 of 12
No
No
2 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
Danio rerio (Zebrafish) (37)
3 of 15
Yes
Yes
3 of 15
Yes
Yes
3 of 15
Yes
Yes
2 of 15
No
No
 
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (11)
2 of 15
Yes
No
 
2 of 15
Yes
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (12)
2 of 9
Yes
Yes
1 of 9
No
No
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
Yes
1 of 9
No
No
1 of 9
No
Yes
1 of 9
No
No
1 of 9
No
Yes
Saccharomyces cerevisiae (Brewer's yeast) (2)
1 of 15
Yes
Yes
1 of 15
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG09190EEV )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG09150914 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( None identified )
No non-Dipteran orthologies identified
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( None identified )
No non-Insect Arthropod orthologies identified
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
No non-Arthropod Metazoa orthologies identified
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (19)
3 of 10
2 of 10
2 of 10
2 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    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
    Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
     
    Disease Associations of Human Orthologs (via DIOPT v7.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    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:
    Select Layout:
    Legend:
    Protein
    RNA
    Selected Interactor(s)
    Interactions Browser

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    esyN Network Diagram
    esyN Network Key:
    Suppression
    Enhancement

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    External Data
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Gene Group - Pathway Membership (FlyBase)
    External Data
    Linkouts
    SignaLink - A signaling pathway resource with multi-layered regulatory networks.
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    3R
    Recombination map
    3-48
    Cytogenetic map
    Sequence location
    3R:6,752,864..6,754,194 [-]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    84A5-84A5
    Limits computationally determined from genome sequence between P{PZ}pb04498 and P{lacW}l(3)L2100L2100
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    84A-84B
    (determined by in situ hybridisation)
    84B1-84B2
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Location
    3-47.5
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (16)
    Genomic Clones (27)
    cDNA Clones (9)
     

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

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

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

      cDNA Clones, End Sequenced (ESTs)
      BDGP DGC clones
        RNAi and Array Information
        Linkouts
        DRSC - Results frm RNAi screens
        GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
        Antibody Information
        Laboratory Generated Antibodies
         
        Commercially Available Antibodies
         
        Other Information
        Relationship to Other Genes
        Source for database identify of
        Source for identity of: zen CG1046
        Source for database merge of
        Additional comments
        Other Comments
        The zen product negatively regulates dac expression.
        dl can activate transcription from zen and twi promoters, and additional Dsp1 inhibits the zen activation and increases the twi activation.
        zen directly activates the amnioserosa-specific expression of a downstream target gene, Ance. zen is an essential, but not sole mediator of the dpp pathway.
        zen ventral repression element (VRE) activity has been studied.
        Binding sites for the grh transcription factor contribute to the ventral repression of zen.
        A putative dpp/zen target gene, Ance, is a homologue of the angiotensin converting enzyme is expressed in the presumptive amnioserosa.
        DNApol-α180 and mus209 are negatively regulated by zen protein. This repression is mediated by the DRE (DNA replication-related element) sites in the DNApol-α180 and mus209 promoters. The amount of Dref is reduced in transfected Kc cells expressing zen, suggesting that zen represses expression of DNA replication-related genes by reducing the amount of Dref.
        A minimal 110bp Ventral Repression Element silencer in the zen promoter contains two dl binding sites as well as binding sites for additional nuclear factors present in early embryos. Mutations in the latter convert the minimal VRE into an enhancer, mediating transcriptional activation in ventral regions in response to dl.
        dl binding sites from the zen promoter can mediate transcriptional activation of a heterologous promoter, but not repression. T-rich sequences close to the dl binding sites in the silencer region of the zen promoter are conserved between D.melanogaster, D.virilis and D.pseudoobscura.pseudoobscura.
        The homologs of Antp, ftz, Scr, Dfd, Ama, bcd, zen, pb and lab, but not zen2 are all present in D.pseudoobscura.pseudoobscura, in the same linear order and similarly spaced along the chromosome as in D.melanogaster.
        Comparative analysis of the homeobox sequences reveals the subdivision of the Antp-type homeobox genes into three classes early in metazoan evolution, one includes Abd-B, the second includes abd-A, Ubx, Antp, Scr, Dfd and ftz, and the third includes zen, zen2, pb and lab.
        zen alleles display relative phenotypic strengths, this may be correlated to the progressive loss of dorsal pattern elements in the ventralised mutants.
        zen expression assayed in embryos from females expressing dl-lacZ fusion proteins compared to wild type dl. Consistant with degree of dorsalization, domain of zen expression is expanded laterally.
        dl binding site domain exchange experiments, using Ecol\lacZ reporter gene constructs, between the zen and twi promoters demonstrate that dl is intrinsically an activator and that repression requires additional factors present in the distal region of the zen promoter, the VR.
        zen gene product is required for proper development of the amnioserosa.
        The zygotically acting DV genes repress ac expression within specific DV domains.
        dl is a sequence specific DNA binding protein that may mediate long range repression by interacting with the distal regions of the zen promoter.
        dl-binding sites present in the twi promoter possess a lower affinity to those present in the zen promoter.
        Mutations in zygotic dorsal class gene zen do not interact with RpII140wimp.
        zen has altered pattern of expression in ventralized and lateralized embryos. In dorsalized embryos zen expression refines at stage 5. Polar expression of zen requires genes of the terminal group. zen is required for the normal ontogeny of the zen pattern and fating of the amnioserosa.
        zen represses transcription from the mus209 promoter of mus209-Ecol\CAT reporter constructs.
        Mutations result in loss of several dorsally derived embryonic structures, disruption of germ band extension and disruption of head involution. Optic lobe and the amnioserosa are most sensitive to loss of zen+ activity. Analysis of a temperature sensitive allele demonstrates the time of zen function is from 2--4 hours of embryogenesis.
        Involved in the regulatory hierarchy responsible for the asymmetric distribution and function of zygotic regulatory gene products along the DV axis of early embryos.
        In vitro DNA footprinting analysis shows that eve and zen can bind to sites in the mus209 5' flanking region.
        The zen promoter shows a two-tier organization: distal sequences mediate its initial response to maternal factors, whereas proximal sequences are responsible for the refinement of the pattern in older embryos. The distal element has the property of a silencer (or anti-enhancer). The proximal element may interact with factors that may be modulated by a cell-cell communication pathway.
        A transient expression assay has been employed to investigate the potential of homeobox genes to function as transcriptional activators.
        The DNA binding activities of the en, eve, prd and zen proteins have been compared.
        P element mediated transformation experiments demonstrate that zen alone can provide zen+ gene function. This suggests that the neighbouring zen2 gene may be dispensable.
        Null mutations result in embryonic lethality and the loss of several dorsally derived embryonic structures, including the amnioserosa, optic lobe and dorsal ridge. These animals also fail to fully extend their germ bands and go through the process of head involution. Hypomorphic mutations result in the absence of dorsal structures but do undergo normal gastrulation movements. A temperature-sensitive allele has been used to define the time of zen+ action between 2 and 4 hours of embryogenesis, just prior to and overlapping the earliest observable morphogenic defects. X-ray induced somatic clones have further shown that zen+ function is unnecessary for postembryonic development. The RNA product of zen is first detected at about 2 hours of development during the eleventh to twelfth cell cycle of the syncytial blastoderm. At this early stage the RNA is found on the dorsal surface of the embryo extending around the anterior and posterior poles. As cellularization proceeds and the early events of gastrulation begin, the RNA becomes restricted to a mid-dorsal stripe of cells. These cells have been fate mapped and give rise to the amnioserosa and the lobes in the dorsal posterior of the embryonic head, i.e., the structures absent in zen- animals. The time of appearance of zen RNA also correlates nicely with the temperature-sensitive-period data obtained using the conditional allele. Antisera to the zen protein product has been used to follow its accumulation pattern and this analysis agrees with and expands the in situ results. The protein is located in the nuclei of cells expressing the gene and at cellular blastoderm is found in a mid-dorsal stripe seven cells wide and seventy cells in length. During gastrulation these cells eventually give rise to the amnioserosa, the optic lobe and dorsal ridge; these structures continue to show zen protein accumulation until the end of germ-band extension at about 4 to 6 hours of development. This end point also correlates well with the end of the temperature-sensitive period of the conditional allele. The spatial pattern of zen expression has been shown to be dependent on the products of several of the maternally expressed genes which specify the anterior-posterior and dorsal-ventral polarity of the embryo, and zen would appear to lie near the end of the axis-determining pathway.
        Origin and Etymology
        Discoverer
        Etymology
        The name for the locus derives from the characteristic 'wrinkled' appearance of the germ band seen in the SEM at the time of normal germ-band retraction.
        Identification
        External Crossreferences and Linkouts ( 35 )
        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/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
        Other crossreferences
        BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
        Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
        Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
        Flygut - An atlas of the Drosophila adult midgut
        GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
        KEGG Genes - Molecular building blocks of life in the genomic space.
        modMine - A data warehouse for the modENCODE project
        SignaLink - A signaling pathway resource with multi-layered regulatory networks.
        Linkouts
        BioGRID - A database of protein and genetic interactions.
        DPiM - Drosophila Protein interaction map
        DroID - A comprehensive database of gene and protein interactions.
        DRSC - Results frm RNAi screens
        FLIGHT - Cell culture data for RNAi and other high-throughput technologies
        FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
        FlyMine - An integrated database for Drosophila genomics
        Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
        InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
        MIST (protein-protein) - An integrated Molecular Interaction Database
        Synonyms and Secondary IDs (17)
        Reported As
        Symbol Synonym
        BG:DS00276.9
        l(3)84Ad
        zen
        (Shokri et al., 2019, Chambers et al., 2017, Jeibmann et al., 2017, Koenecke et al., 2017, Transgenic RNAi Project members, 2017-, Bürglin and Affolter, 2016, Quijano et al., 2016, Sandler and Stathopoulos, 2016, Sarov et al., 2016, Schmidt-Ott and Kwan, 2016, O'Connell and Reeves, 2015, Ugrankar et al., 2015, Künnapuu et al., 2014, Mannervik, 2014, Shin and Hong, 2014, Chen et al., 2013, Heffer and Pick, 2013, Saunders et al., 2013, Upadhyai and Campbell, 2013, Holmqvist et al., 2012, Kim et al., 2012, Kvon et al., 2012, Liang et al., 2012, Reeves et al., 2012, Turki-Judeh and Courey, 2012, Chung et al., 2011, Garcia and Stathopoulos, 2011, Gehring, 2011, Harrison et al., 2011, Kim et al., 2011, Mrinal et al., 2011, Nien et al., 2011, Ozdemir et al., 2011, Roy et al., 2011, Tsurumi et al., 2011, Wilson and Dearden, 2011, Bothma et al., 2010, Harrison et al., 2010, Hueber et al., 2010, Kazemian et al., 2010, Rafiqi, 2010.5.12, Salzer et al., 2010, Su et al., 2010, Tipping et al., 2010, Bai et al., 2009, Christensen et al., 2008.4.15, Christensen et al., 2008.4.15, Ishihara and Shibata, 2008, Kwong et al., 2008, Liang et al., 2008, Li et al., 2008, Miles et al., 2008, Pope and Harris, 2008, Aerts et al., 2007, Astigarraga et al., 2007, Beltran et al., 2007, Copley et al., 2007, Demuth and Wade, 2007, Duboule, 2007, Gunaydin et al., 2007, Kumar and Anderson, 2007, Negre and Ruiz, 2007, Nien et al., 2007, Roy et al., 2007, Ullah et al., 2007, Zeitlinger et al., 2007, Brandt, 2006, Casillas et al., 2006, Chen et al., 2006, Friedrich, 2006, Lin et al., 2006, Ratnaparkhi et al., 2006, Scuderi et al., 2006, Shirangi et al., 2006, ten Bosch et al., 2006, Barker et al., 2005, Hoskins et al., 2005, Negre et al., 2005, Pearson et al., 2005, Peel et al., 2005, Schmidt-Ott, 2005, Stathopoulos and Levine, 2005, Gurunathan et al., 2004, Riede, 2004, Jia et al., 2002, Kazemi-Esfarjani and Benzer, 2002, Chen et al., 2000, Kidd, 1992)
        Secondary FlyBase IDs
        • FBgn0013967
        • FBgn0019915
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        References (397)