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General Information
Symbol
Dmel\skd
Species
D. melanogaster
Name
skuld
Annotation Symbol
CG9936
Feature Type
FlyBase ID
FBgn0003415
Gene Model Status
Stock Availability
Gene Summary
skuld (skd) encodes a subunit of the kinase module of the mediator complex. It is not required for all the transcriptional functions of mediator, but links it to a set of transcription factors that are involved in developmental signaling. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
Also Known As

pap, Med13, bli, blind spot, flytrap

Key Links
Genomic Location
Cytogenetic map
Sequence location
3L:20,992,814..21,027,434 [-]
Recombination map
3-47
RefSeq locus
NT_037436 REGION:20992814..21027434
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (16 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from physical interaction with UniProtKB:Q9VW47
(assigned by UniProt )
inferred from physical interaction with UniProtKB:P25008
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from sequence or structural similarity with HGNC:22474
inferred from biological aspect of ancestor with PANTHER:PTN000310508
(assigned by GO_Central )
Biological Process (10 terms)
Terms Based on Experimental Evidence (9 terms)
CV Term
Evidence
References
inferred from genetic interaction with FLYBASE:Scr; FB:FBgn0003339
involved_in chaeta development
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from expression pattern
inferred from mutant phenotype
inferred from direct assay
(assigned by UniProt )
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:Scr; FB:FBgn0003339
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000310508
(assigned by GO_Central )
inferred from sequence or structural similarity with HGNC:22474
Cellular Component (4 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:MED17; FB:FBgn0038578
inferred from physical interaction with FLYBASE:CycC; FB:FBgn0004597, FLYBASE:Cdk8; FB:FBgn0015618
inferred from direct assay
(assigned by UniProt )
located_in nucleoplasm
inferred from high throughput direct assay
located_in nucleus
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from sequence or structural similarity with HGNC:22474
inferred from biological aspect of ancestor with PANTHER:PTN000310508
(assigned by GO_Central )
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the Mediator complex subunit 13 family. (Q7KTX8)
Summaries
Gene Snapshot
skuld (skd) encodes a subunit of the kinase module of the mediator complex. It is not required for all the transcriptional functions of mediator, but links it to a set of transcription factors that are involved in developmental signaling. [Date last reviewed: 2019-03-14]
Gene Group (FlyBase)
MEDIATOR COMPLEX -
The Mediator (MED) complex serves as a hub for transcriptional signaling events. MED bridges transcriptional activators and repressors. This large multi-protein complex has a central core conserved from yeast to humans. The core is composed of three modules: head, middle, and tail. A separate module, the Cdk8 kinase module, composed of Cdk8, CycC, kto and skd, serves as a regulator. (Adapted from FBrf0226415 and FBrf0191850).
Pathway (FlyBase)
Positive Regulators of Wnt-TCF Signaling Pathway -
Positive regulators of Wnt-TCF (canonical Wnt) signaling up-regulate the pathway, resulting in the enhancement of transcriptional regulation mediated by β-catenin (arm).
Protein Function (UniProtKB)
Component of the Mediator complex, a coactivator involved in the regulated transcription of nearly all RNA polymerase II-dependent genes. Mediator functions as a bridge to convey information from gene-specific regulatory proteins to the basal RNA polymerase II transcription machinery. Mediator is recruited to promoters by direct interactions with regulatory proteins and serves as a scaffold for the assembly of a functional preinitiation complex with RNA polymerase II and the general transcription factors (By similarity). Required for leg and eye development and macrochaete specification or differentiation. Negatively regulates sex comb development. Required for activated transcription of the MtnB and MtnD genes.
(UniProt, Q7KTX8)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
skd: skuld (J.A. Kennison)
Isolated as a dominant suppressor of Pc mutations. Associated with recessive lethality at the larval-pupal transition. Also interacts with Pcl, Scr, and Ubx mutations. May weakly suppress AntpNs.
Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
3

Please see the JBrowse view of Dmel\skd 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

Annotated transcripts do not represent all possible combinations of alternative exons and/or alternative promoters.

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.46

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0078329
11414
2618
FBtr0078328
10465
2618
FBtr0112830
10703
2768
FBtr0333284
9982
2601
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
FBpp0077985
280.0
2618
6.54
FBpp0077984
280.0
2618
6.54
FBpp0111743
295.7
2768
6.82
FBpp0305479
278.2
2601
6.53
Polypeptides with Identical Sequences

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

2618 aa isoforms: skd-PC, skd-PD
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

Component of the Cdk8 module of the Mediator complex, composed of CycC, Cdk8, kto and skd.

(UniProt, Q7KTX8)
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\skd 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
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
organism

Comment: maternally deposited

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

Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:MED17; FB:FBgn0038578
inferred from physical interaction with FLYBASE:CycC; FB:FBgn0004597, FLYBASE:Cdk8; FB:FBgn0015618
inferred from direct assay
(assigned by UniProt )
located_in nucleoplasm
inferred from high throughput direct assay
located_in nucleus
inferred from direct assay
Expression Deduced from Reporters
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\skd 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
EMBL-EBI Single Cell Expression Atlas
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 46 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 8 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of skd
Transgenic constructs containing regulatory region of skd
Aberrations (Deficiencies and Duplications) ( 6 )
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
15 of 15
Yes
Yes
12 of 15
No
Yes
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (2)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
15 of 15
Yes
Yes
12 of 15
No
Yes
Rattus norvegicus (Norway rat) (2)
13 of 13
Yes
Yes
5 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (3)
9 of 12
Yes
Yes
8 of 12
No
Yes
1 of 12
No
Yes
Danio rerio (Zebrafish) (2)
13 of 15
Yes
Yes
10 of 15
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (1)
12 of 15
Yes
Yes
Arabidopsis thaliana (thale-cress) (1)
2 of 9
Yes
Yes
Saccharomyces cerevisiae (Brewer's yeast) (1)
1 of 15
Yes
No
Schizosaccharomyces pombe (Fission yeast) (1)
1 of 12
Yes
Yes
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG09190074 )
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) ( EOG0915006K )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Lucilia cuprina
Australian sheep blowfly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Anopheles darlingi
American malaria mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W00R7 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X00PR )
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
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G00AP )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (2)
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 ( 1 )
Allele
Disease
Evidence
References
Potential Models Based on Orthology ( 1 )
Modifiers Based on Experimental Evidence ( 2 )
Disease Associations of Human Orthologs (via DIOPT v8.0 and OMIM)
Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
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
Subunit Structure (UniProtKB)
Component of the Cdk8 module of the Mediator complex, composed of CycC, Cdk8, kto and skd.
(UniProt, Q7KTX8 )
Linkouts
BioGRID - A database of protein and genetic interactions.
DroID - A comprehensive database of gene and protein interactions.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Signaling Pathways (FlyBase)
Positive Regulators of Wnt-TCF Signaling Pathway -
Positive regulators of Wnt-TCF (canonical Wnt) signaling up-regulate the pathway, resulting in the enhancement of transcriptional regulation mediated by β-catenin (arm).
Metabolic Pathways
External Data
Linkouts
Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
Genomic Location and Detailed Mapping Data
Chromosome (arm)
3L
Recombination map
3-47
Cytogenetic map
Sequence location
3L:20,992,814..21,027,434 [-]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
78A2-78A2
Limits computationally determined from genome sequence between P{EP}fngEP3082&P{lacW}skdL7062 and P{lacW}l(3)L5541L5541
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
78A1-78A2
(determined by in situ hybridisation) 78A2--3 (determined by in situ hybridisation)
78A-78A
(determined by in situ hybridisation)
78A1-78A2
(determined by in situ hybridisation) 78A2--3
Experimentally Determined Recombination Data
Left of (cM)
Right of (cM)
Notes

Shows low recombination frequency with Ki mutants.

Stocks and Reagents
Stocks (24)
Genomic Clones (30)
cDNA Clones (49)
 

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 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)
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

Source for merge of: pap CG9936

Source for merge of: pap l(3)L7062

Source for merge of: pap bli l(3)rK760

Source for merge of: skd pap

Source for merge of: pap flytrap

Additional comments

snoRNA:skd-a is encoded in an intron of skd.

Source for merge of pap flytrap was sequence comparison ( date:000315 ).

Other Comments

dsRNA has been made from templates generated with primers directed against this gene.

Both skd and kto are required to establish normal cell affinity differences at the anterior-posterior and dorsal-ventral compartment boundaries of the wing disc.

skd and kto seem to act together to promote specific cell fate decisions, transmitting either a novel signal or a combination of signals.

skd is required for the normal progression of photoreceptor differentiation in the eye disc and for the normal patterning of other imaginal discs.

Loss of skd function appears to arrest cells in the eye disc at an early stage of their differentiation.

skd is required for viability.

P-element-induced mutations have been isolated as dominant modifiers in combination with pb gain-of-function transgenes (HSPB). These modifiers are selected by their effects on specific cell identities sensitive to Ras85D signaling activities. One modifier, skd, favors the appearance of sex comb teeth on the more distal second tarsal segment of the T1 leg. skd also modifies the action of pb hypomorphs in mouthparts development, favoring the appearance of sex comb teeth on the adult labial palps. Animals lacking zygotic function die at the end of embryogenesis with mouthparts defects. Hypomorphic alleles from imprecise P excisions show abdominal defects and female sterility.

Sections of the Scr regulatory region may be important for regulation of Scr by Polycomb- and trithorax-group genes.

skd is one of the 18 loci identified in a screen for dominant modifiers of Pc and/or Antp phenotypes. Alleles of Pc, Pcl, Scm, Dll, brm, kto, Scr and trx show clear dominant enhancement or suppression of AntpScx, whereas alleles of vtd, Vha55, Su(Pc)37D, urd, mor, skd and osa do not.

Origin and Etymology
Discoverer

Kennison, 1984.

Etymology

'poils aux pattes' is French for 'hairy legs'.

Identification
External Crossreferences and Linkouts ( 69 )
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 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
EMBL-EBI Single Cell Expression Atlas
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
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)
KEGG Genes - Molecular building blocks of life in the genomic space.
MARRVEL_MODEL
modMine - A data warehouse for the modENCODE project
Linkouts
BioGRID - A database of protein and genetic interactions.
DroID - A comprehensive database of gene and protein interactions.
DRSC - Results frm RNAi screens
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
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
Synonyms and Secondary IDs (36)
Reported As
Symbol Synonym
Srb9/AMIB/PAP/TRAP240
l(3)L7062
pap/dTRAP240
Secondary FlyBase IDs
  • FBgn0024200
  • FBgn0024569
  • FBgn0024577
  • FBgn0037043
  • FBgn0041162
Datasets (0)
Study focus (0)
Experimental Role
Project
Project Type
Title
References (156)