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General Information
Symbol
Dmel\ems
Species
D. melanogaster
Name
empty spiracles
Annotation Symbol
CG2988
Feature Type
FlyBase ID
FBgn0000576
Gene Model Status
Stock Availability
Gene Snapshot
empty spiracles (ems) encodes a homeotic selector gene that encodes a transcription factor involved in spiracle morphogenesis, axogenesis, and development of the ventral nerve cord, brain and tracheal system. [Date last reviewed: 2019-03-07]
Key Links
Genomic Location
Cytogenetic map
Sequence location
3R:13,901,853..13,904,620 [+]
Recombination map

3-54

RefSeq locus
NT_033777 REGION:13901853..13904620
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
Belongs to the EMX homeobox family. (P18488)
Summaries
Gene Group (FlyBase)
NK-LIKE HOMEOBOX TRANSCRIPTION FACTORS -
NK-like (NKL) homeobox transcription factors are sequence-specific DNA binding proteins that regulate transcription. NKL transcription factors are homeobox genes closely related to Hox-like genes, a number of which are found in the NK cluster. Many of the NKL members contain an Engrailed Homology 1 (EH1) motif. (Adapted from FBrf0232555 and PMID:22094586).
Protein Function (UniProtKB)
Acts as a homeotic selector gene controlling antennal and mandibular segment identity.
(UniProt, P18488)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
ems: empty spiracles
Embryonic lethal. Embryos display loss of filzorper in the posterior spiracles; posterior ends of longitudinal tracheal trunks incomplete. Failure of head involution; many cuticular structures that normally derive from the procephalic and mandibular lobes of the head missing; embryonic antennal organs missing. ems protein first appears as an anterior ring around the syncytial blastoderm at a position just anterior to that of Dfd; it is five to six cells wide dorsally and 10-12 cells wide ventrally; at gastrulation the ring is just anterior to the cephalic furrow. As gastrulation proceeds expression becomes patchy and confined to specific groups of cells. The protein is nuclear in localization. The metameric pattern of ems expression commences at the beginning of germ-band extension; protein first appears in a group of cells in each segment, which subsequently elongates and splits into two clusters of ems-positive cells in regions destined to give rise to tracheal pits, neuroblasts and epidermis. In the eighth abdominal segment, a large patch of ems-positive cells forms just posterior to the tracheal pit and presumably correspond to the primordia of the posterior spiracles and the filzkorper; a similar patch is formed anteriorly in embryos produced by bcd mothers. For a detailed discussion of the expression pattern see Dalton et al..
Summary (Interactive Fly)

transcription factor - homeodomain - required for head development and brain morphogenesis - required for the development of olfactory projection neuron circuitry

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

Please see the GBrowse view of Dmel\ems or the JBrowse view of Dmel\ems 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 5.53

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0082863
2478
494
Additional Transcript Data and Comments
Reported size (kB)

2.4 (northern blot)

2.3 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0082326
53.4
494
7.60
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)

497 (aa); 54 (kD predicted)

Comments
External Data
Crossreferences
InterPro - A database of protein families, domains and functional sites
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\ems 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 (19 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000678271
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000678271
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000678271
(assigned by GO_Central )
Biological Process (14 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from expression pattern
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
traceable author statement
non-traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000678271
(assigned by GO_Central )
non-traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000678271
(assigned by GO_Central )
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000678271
(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
antennal anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

dorsal head epidermis 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

antennal primordium

Comment: reported as procephalic ectoderm primordium

central brain primordium

Comment: reported as procephalic ectoderm primordium

visual primordium

Comment: reported as procephalic ectoderm primordium

dorsal head epidermis primordium

Comment: reported as procephalic ectoderm primordium

lateral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

ventral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

dorsal epidermis primordium

Comment: reported as dorsal epidermis anlage

gnathal primordium

Comment: reported as gnathal lobes anlage

tracheal primordium

Comment: reported as tracheal system anlage

organism | 70-90% egg length

Comment: 70-76% egg length dorsally; 74-89% ventrally

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

ems transcripts are expressed at roughly 69-75% egg length in the cellular blastoderm embryo. Beginning at gastrulation, they are expressed in a set of stripes just anterior to the cephalic furrow. Expression of ems in the protocerebral domain of the head is mostly gone by stage 9.

ems transcripts are detected at all developmental stages except adult on northern blots. The peak of expression occurs between 6 and 12 hours of embryogenesis and a second smaller peak is seen in second instar larvae. Transcripts are first detected by in situ hybridization at the syncytial blastoderm stage in a stripe that extends from 70-76% egg length dorsally and 74-89% egg length ventrally. At the cellular blastoderm stage, expression fades in the ventral part of the stripe and to a lesser extent dorsally. At stages 6 and 7, @ems transcripts are expressed anterior to and along the cephalic furrow and in the anlagen for the intercalary and antennal segments and in part of the procephalic lobe. At stage 8, they are observed in lateral regions along the extending germ band and in an anterior dorsal patch at 85-90% egg length. During stage 9, expression is seen in 16 regions along the extended germ band and in the mandibular, intercalary, and antennal segments. By stage 10, staining in the gnathal segments corresponds to the anterior part of each segment and the staining in the abdominal segments occurs around the tracheal pits. Staining is also observed in the primordia of the posterior spiracles. Staining in the optic lobe primordium begins at stage 12. By stage 13, expression is seen in lateral cells of the CNS and in a row of cells immediately posterior to the forming segmental furrows. In stage 14, expression is restricted to the anterior part of the mandibular lobe, the antenno-maxillary complex, the subesophageal ganglion, and the posterior spiracles

ems transcripts are expressed in all stages tested with a peak in 6-12 hr embryos, and high levels in 3rd instar larvae and pupae.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
cephalic furrow | anterior to

Comment: single circumferential stripe of expression

Additional Descriptive Data

At 0 hr APF, ems is detected in superficial cells located along the periphery of the antennal disc, in the second antennal segment, and at the base of the presumptive arista probably within the epidermal cells of the disc. In the third antennal segment of the disc, initial ems expression is observed at 0 hr APF, and by 2 hr APF this increases to include cells in four semi-circular domains. Expression in these domains broadens by 5 hours APF. ems levels diminish by 8 hr APF and it is undetectable by 12 hr APF. ems is present in a subset of amos-expressing cells but does not overlap ato expression. The time course of onset is similar to amos but it is down regulated earlier. ems is also expressed in a subset of lz-expressing cells and transiently in a subset of the sens-expressing proneural domains from which SOPs are selected. In summary, ems is expressed, together with amos, in a subset of proneural clusters in the third segment of the antennal disc. Its expression is transient in SOPs that are selected from these clusters indicating that it may be involved in the development of a subset of the trochoid and/or bas iconic olfactory sense organs.

ems protein is detected in discrete domains in the procephalic ectoderm of stage 9-11 embryos, including initially 3 domains between the anterior part of the mandibular segment and the posterior of the ocular segment. One of these domains further splits. Additional small domains of expression are observed in another region of the ocular segment and in a dorsal patch of procephalic ectoderm. Expression in procephalic neuroblasts stage 9-11: tritocerebrum - d6; deuterocerebrum - d3, d6, v3, v7; protocerebrum - cv5, pd2, pd5, pd8, pd10, pd15

In stage 6 embryos, the ems protein expression domain is posterior to that of tll, with a 1-2 cell unlabled gap between. By embryonic stage 10, ems protein is distributed in a segmentally striped pattern. At this stage, ems expression colocalizes with tll in the intercalary segment, and is adjacent to tll in the antennal and ocular segments. At stage 15, ems expression is adjacent to tll expression in the embryonic brain.

ems is expressed in two independently regulated patterns that are described in detail. The head specific pattern initiates prior to cellular blastoderm and continues to early germband extension. The metameric pattern initiates after gastrulation and includes expression in lateral neuroblasts, in ectodermal cells at the anterior lateral borders of each segment and in the filzkorper primordia. The region of ems expression in the head is anterior and adjacent to the Dfd expression stripe. In bcd mutants, the blastoderm head specific ems staining is absent while the later metameric pattern is unaffected.

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

GBrowse - Visual display of RNA-Seq signals

View Dmel\ems 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
Alleles, Insertions, and Transgenic Constructs
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (23)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
6 of 15
Yes
No
5 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
Yes
1 of 15
No
No
1 of 15
No
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (26)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
6 of 15
Yes
No
 
 
5 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
Yes
1 of 15
No
No
 
1 of 15
No
No
Rattus norvegicus (Norway rat) (21)
6 of 13
Yes
No
5 of 13
No
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
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
Xenopus tropicalis (Western clawed frog) (22)
5 of 12
Yes
No
5 of 12
Yes
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
Yes
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
Yes
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) (28)
6 of 15
Yes
No
5 of 15
No
No
4 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
Yes
1 of 15
No
No
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (11)
6 of 15
Yes
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
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
Arabidopsis thaliana (thale-cress) (1)
1 of 9
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG09190DVB )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
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 grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG09150AAM )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0CM5 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0CK6 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0P41 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (15)
4 of 10
2 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
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
    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.
    Dmel gene
    Ortholog showing functional complementation
    Supporting References
    Interactions
    Summary of Physical Interactions
    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
    DroID - A comprehensive database of gene and protein interactions.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    External Data
    Linkouts
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    3R
    Recombination map

    3-54

    Cytogenetic map
    Sequence location
    3R:13,901,853..13,904,620 [+]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    88A2-88A2
    Limits computationally determined from genome sequence between P{PZ}flfl01949 and P{lacW}trxj14A6&P{PZ}trx00347
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    88A1-88A2
    (determined by in situ hybridisation)
    88A-88A
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (13)
    Genomic Clones (17)
     

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

    cDNA Clones (16)
     

    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 database merge of
      Additional comments
      Other Comments

      ems is essential for correct post-embryonic projection neuron development in the olfactory system.

      In the posterior brain and the ventral nerve cord, ems is necessary for correct axonal pathfinding of specific interneurons.

      Mis-expression of ems in the head transforms segment identity in a btd-dependent manner.

      ems and btd gene products interact in vitro.

      ems is able to specify head identity when acting in concert with btd.

      In ems mutants, invagination of the spiracle cells adjacent to the trachea does not occur, though more posterior cells of the spiracle invaginate normally, producing a spiracle without a lumen.

      In a sample of 79 genes with multiple introns, 33 showed significant heterogeneity in G+C content among introns of the same gene and significant positive correspondence between the intron and the third codon position G+C content within genes. These results are consistent with selection adding against preferred codons at the start of genes.

      Psc protein is associated with an upstream fragment of ems.

      Probes labelled with digoxigenin, fluorescein and biotin allow detection of RNA of three different genes in three different colours.

      ems is the gene responsible for sclerotic plate formation in larvae lacking all thoracic and abdominal homeotic genes and is phenotypically suppressed by the homeotic genes in normal development.

      oc and ems are required for brain segmentation in the embryo.

      Heat shock induced expression of mouse Hox genes in Drosophila embryos deficient for homeotic genes demonstrates that functional hierarchy is a universal property of the homeobox genes. Correlations exist between the expression patterns of the mouse Hox genes along the antero-posterior body axis of mice and the extent of their effect along the antero-posterior body axis of flies.

      ems, a head-specific gap gene, may function as a co-repressor of dl, thereby linking the anteroposterior and dorsoventral systems st the molecular level.

      Expression of the trn pair rule stripes requires ftz and ems.

      ems is a downstream gene that is transcriptionally regulated by Abd-B gene products. Genetic epistasis tests suggest that both ems and Abd-B gene products are required in combination for the specification of the filzkorper primordia.

      One of the homeodomain loci identified in a screen for genes encoding DNA binding proteins capable of binding to a consensus Engrailed binding site.

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

      Isolated by cross-homology to a Dr probe. Stated to be at 88A: no further details given.

      ems has been cloned and sequenced and shown to be a homeobox-containing gene. Its expression pattern and mutant phenotype suggest it is a gap gene.

      ems has a gap-gene-like role in mediating the development of head structures. In ems mutant embryos gt stripe 1 is normal but stripes 1 and 2 fail to sharpen and separate at cellular blastoderm. ems is positively regulated by bcd.

      Mutations in zygotic gene ems do not interact with RpII140wimp.

      btd, ems and oc are required to establish contiguous blocks of segments and may behave like gap genes that mediate bcd function in the embryonic head.

      ems is required for the development of structures that derive from the most anterior head segments of the embryo.

      ems mutants display spiracles devoid of filzkorper, no antenna and an open head.

      Embryonic lethal. Embryos display loss of filzkorper in the posterior spiracles; posterior ends of longitudinal tracheal trunks incomplete. Failure of head involution; many cuticular structures that normally derive from the procephalic and mandibular lobes of the head missing; embryonic antennal organs missing. ems protein first appears as an anterior ring around the syncytial blastoderm at a position just anterior to that of Dfd; it is five to six cells wide dorsally and 10-12 cells wide ventrally; at gastrulation the ring is just anterior to the cephalic furrow. As gastrulation proceeds expression becomes patchy and confined to specific groups of cells. The protein is nuclear in localization. The metameric pattern of ems expression commences at the beginning of germ-band extension; protein first appears in a group of cells in each segment, which subsequently elongates and splits into two clusters of ems-positive cells in regions destined to give rise to tracheal pits, neuroblasts and epidermis. In the eighth abdominal segment, a large patch of ems-positive cells forms just posterior to the tracheal pit and presumably correspond to the primordia of the posterior spiracles and the filzkorper; a similar patch is formed anteriorly in embryos produced by bcd mothers. For a detailed discussion of the expression pattern see Dalton et al.

      Origin and Etymology
      Discoverer
      Etymology
      Identification
      External Crossreferences and Linkouts ( 33 )
      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
      UniProt/TrEMBL - Automatically annotated and unreviewed 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
      iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
      InterPro - A database of protein families, domains and functional sites
      KEGG Genes - Molecular building blocks of life in the genomic space.
      modMine - A data warehouse for the modENCODE project
      Linkouts
      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
      FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
      FlyMine - An integrated database for Drosophila genomics
      Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
      MIST (protein-protein) - An integrated Molecular Interaction Database
      Synonyms and Secondary IDs (14)
      Reported As
      Symbol Synonym
      W13
      ant
      ems
      (Banerjee et al., 2019, Cai et al., 2019, Curt et al., 2019, Shokri et al., 2019, Steinmetz et al., 2019, Bischof et al., 2018, Burow et al., 2018, Wang et al., 2018, Hu et al., 2017.6.13, Karaiskos et al., 2017, Transgenic RNAi Project members, 2017-, Bürglin and Affolter, 2016, Hauptmann et al., 2016, Peng et al., 2016, Skottheim Honn et al., 2016, Urbach et al., 2016, Enriquez et al., 2015, Glassford et al., 2015, Schertel et al., 2015, Palsson et al., 2014, Sen et al., 2014, Spahn et al., 2014, Tsikala et al., 2014, Li and Gilmour, 2013, Matzat et al., 2013, Saunders et al., 2013, Sen et al., 2013, Webber et al., 2013, Andrioli et al., 2012, Chen et al., 2012, He et al., 2012, Immonen and Ritchie, 2012, Nikulova et al., 2012, Bellen et al., 2011, Gehring, 2011, Perry et al., 2011, Zhai et al., 2011, Hartmann et al., 2010, Kazemian et al., 2010, Maurel-Zaffran et al., 2010, Sen et al., 2010, Yassin et al., 2010, Zhai et al., 2010, Kumar et al., 2009, Löhr et al., 2009, Nanda et al., 2009, Ochoa-Espinosa et al., 2009, Seibert et al., 2009, Stöbe et al., 2009, Venken et al., 2009, Venken et al., 2009, Blanco and Gehring, 2008, Das et al., 2008, DeFalco et al., 2008, Lichtneckert et al., 2008, Sotillos et al., 2008, Hueber et al., 2007, Krattinger et al., 2007, Lichtneckert et al., 2007, Magalhaes et al., 2007, Sprecher et al., 2007, Lovegrove et al., 2006, Lovegrove et al., 2006, Scholtz and Edgecombe, 2006, Sprecher and Hirth, 2006, Urbach et al., 2006, Copley, 2005, Grad et al., 2004, Hauptmann, 2001)
      Secondary FlyBase IDs
      • FBgn0000145
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      References (287)