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General Information
Symbol
Dmel\tud
Species
D. melanogaster
Name
tudor
Annotation Symbol
CG9450
Feature Type
FlyBase ID
FBgn0003891
Gene Model Status
Stock Availability
Gene Snapshot
In progress.Contributions welcome.
Key Links
Genomic Location
Cytogenetic map
Sequence location
2R:21,183,322..21,192,913 [+]
Recombination map
2-93
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
-
Predictions / Assertions
-
Summaries
Protein Function (UniProtKB)
Required during oogenesis for the formation of primordial germ cells and for normal abdominal segmentation.
(UniProt, P25823)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
tud: tudor (T. Schupbach)
exhibit a so-called "grandchildless-knirps" phenotype: all eggs lack polar granules and no pole cells are formed; most of the embryos show variable deletions of abdominal segments, whereby segment A4 is deleted most frequently; larger deletions may include segments A2 through A7; in extreme cases anterior parts of segment A1 become fused to posterior part of segment A8, but telson elements are always present and relatively normal. Around 30% of all embryos survive and grow into sterile adults. Analysis of germline clones indicates that the mutation is germline autonomous (Schupbach and Wieschaus, 1986, Dev. Biol. 113: 443-48).
Summary (Interactive Fly)
tudor domain protein - serves as 'docking platform' for polar granule assembly - maternal effect gene required for germ cell formation and abdominal segmentation during oogenesis - interacts with Aubergine, a Piwi family protein, in a manner dependent on symmetrically dimethylated arginine residues located at the N-terminal end of Aub
Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
2

Please see the GBrowse view of Dmel\tud or the JBrowse view of Dmel\tud 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
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.51
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0071582
8830
2515
FBtr0342870
8821
2512
Additional Transcript Data and Comments
Reported size (kB)
8.0 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0071508
285.3
2515
6.40
FBpp0309672
284.9
2512
6.37
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)
2515 (aa); 285 (kD observed)
2515 (aa); 285 (kD predicted)
Comments
External Data
Subunit Structure (UniProtKB)
Interacts with vls (PubMed:15800004). Interacts with me31B (when dimethylated on Arg residues) (PubMed:28945271).
(UniProt, P25823)
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\tud 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 (10 terms)
Molecular Function (0 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (5 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
Cellular Component (5 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
inferred from direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
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
Additional Descriptive Data
Northern analysis indicates that the 8 kb tud transcript is present in all stages of development. During oogenesis, tud transcript is first detected in germarium region 2, in one cell within each cyst. This cell occupies a posterior position in later cysts, and is likely to be the oocyte precursor. In stage S1 to S2 egg chambers, tud transcript is detected completely surrounding the oocyte nucleus. From stage S4 to S7 of oogenesis, the tud transcript is detected in the posterior-most region of the oocyte. The amount of tud transcript detected decreases until stage S8, after which no signal is detected in the oocyte. From stage S10 to S14, nurse cells produce large amounts of tud transcript, which is apparently only transiently present in the oocyte, as no tud transcript is detected by in situ localization in late oogenesis, at egg deposition, or through early embryogenesis.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
tud protein is detected in the germarium in early oogenesis. By stage S4-S6, tud protein is at the anterior margin of the oocyte, and by stage S6, tud protein is detected at the posterior portion of the oocyte. At stage S10 of oogenesis, tud protein is located in a thin crescent at the posterior of the oocyte. This posterior localization is seen in the unfertilized egg, as well as during the early embryonic stages up to cellularization. EM immunolocalization studies indicate that tud protein is present in the posterior pole plasm, with some of the protein localized to the polar granules, as well as in the mitochondria of cells throughout the early embryo.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
inferred from direct assay
inferred from direct assay
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\tud 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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
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, and Transgenic Constructs
Classical and Insertion Alleles ( 35 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 22 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of tud
Transgenic constructs containing regulatory region of tud
Deletions and Duplications ( 17 )
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 v7.1)
Homo sapiens (Human) (7)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
6 of 15
Yes
Yes
4 of 15
No
Yes
2 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (5)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
5 of 15
Yes
Yes
4 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Rattus norvegicus (Norway rat) (6)
2 of 13
Yes
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) (2)
2 of 12
Yes
Yes
1 of 12
No
No
Danio rerio (Zebrafish) (9)
3 of 15
Yes
No
2 of 15
No
No
2 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
Caenorhabditis elegans (Nematode, roundworm) (4)
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
Yes
Arabidopsis thaliana (thale-cress) (0)
No records found.
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) ( EOG09190057 )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
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) ( EOG0915018J )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W03ZA )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble 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
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X00ZN )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
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
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G01J3 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (10)
3 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
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
Subunit Structure (UniProtKB)
Interacts with vls (PubMed:15800004). Interacts with me31B (when dimethylated on Arg residues) (PubMed:28945271).
(UniProt, P25823 )
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
Genomic Location and Detailed Mapping Data
Chromosome (arm)
2R
Recombination map
2-93
Cytogenetic map
Sequence location
2R:21,183,322..21,192,913 [+]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
57C8-57C9
Limits computationally determined from genome sequence between P{EP}CG4266EP2258 and P{EP}CG30394EP962&P{lacW}domk08108
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
57C-57C
(determined by in situ hybridisation)
57C7-57C8
(determined by in situ hybridisation)
Experimentally Determined Recombination Data
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (16)
Genomic Clones (14)
 

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

cDNA Clones (42)
 

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
        Other 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
          Eight alleles, l(2)57Ce1 to l(2)57Ce8, complement both lethal and grandchildless phenotypes of tud (O'Donnell et al., 1989).
          Other Comments
          tud is essential for pole cell specification and polar granule formation but is dispensible for somatic posterior patterning and oogenesis.
          tud may mediate the transport of mitochondrial rRNAs from mitochondria to polar granules.
          tud contains a Trf-responsive promoter.
          osk, stau, vas and tud are essential for pole plasm formation. vas and tud are localised dependent of osk protein and are required to accumulate osk protein stably at the posterior pole.
          tud males produce seminal fluid but no spermatozoa. Lifespan of females mated to Cbβ\DT-A males is greater than those mated to tud males. The entire cost of mating is attributable to receiving male main-cell products, the main cell products have a quantitative effect. The main-cell products are involved in sperm competition, tud males incapacitated 80% of sperm stored by females during a previous mating, males thus increase the proportion of offspring they sire.
          Mutations in tud affect two distinct determinative processes in embryogenesis: segmentation in the abdomen and determination of the primordial germ cells. The tud protein is concentrated in the posterior pole cytoplasm where it is found in polar granules and mitochondria. Throughout the rest of the embryo it is associated with cleavage nuclei. Mutations that affect the germ plasm eliminate the posterior localization, and mutations that affect segmentation disrupt the localization around the nuclei.
          There is no cost to the female to receive sperm: lifespan, egg production, egg hatchability and remating rate of females intermittently exposed to males that could (tra mutants) or could not (tud mutants) transfer sperm are not significantly different.
          Only vas and tud are essential for osk-induced pole cell and abdomen formation.
          vas, vls and tud (but not stau, capu, or spir) are necessary for the 6xosk mutant phenotypes.
          tud is not required for the nurse cell synthesis of the posterior signal.
          Mutations in maternal posterior class gene tud do not interact with RpII140wimp.
          tud plays a role in early embryogenesis and, perhaps, in pattern formation.
          Grandchildless-knirps class of genes.
          tud plays a role in polar granule formation.
          Identification: Transcription unit identified during molecular analysis of the Pu gene region.
          Mutation in tud results in a maternal effect "grandchildless knirps-like" phenotype.
          Mutations in tud cause failure of germ cell formation and deletions in the abdominal segments in the embryo.
          Origin and Etymology
          Discoverer
          Wieschaus and Nusslein-Volhard.
          Etymology
          Identification
          External Crossreferences and Linkouts ( 38 )
          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
          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.
          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)
          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
          BioGRID - A database of protein and genetic interactions.
          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 (9)
          Reported As
          Symbol Synonym
          R5_2R:17070527..17070890
          Secondary FlyBase IDs
            Datasets (0)
            Study focus (0)
            Experimental Role
            Project
            Project Type
            Title
            References (323)