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Gene Dmel\tud

General Information
Symbol	Dmel\tud	Species	D. melanogaster
Name	tudor	Annotation symbol	CG9450
Feature type	protein_coding_gene	FlyBase ID	FBgn0003891
Created / Updated	2003-12-01/2003-12-01
Genomic Location
Chromosome (arm)	2R	Recombination map
Cytogenetic map	57C8-57C9	Sequence location	2R:17,070,937..17,079,903 [+]
Map ( GBrowse )	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene tudor is referred to in FlyBase by the symbol tud (CG9450, FBgn0003891). It has the cytological map location 57C8-57C9. Its sequence location is 2R:17070937..17079903. Its molecular function is described as nucleic acid binding. It is involved in the biological processes: pole plasm assembly; embryonic development; mitochondrial rRNA export from mitochondrion; intracellular mRNA localization; pole cell fate determination; germ cell development; polar granule organization and biogenesis; pole cell development. 38 alleles are reported. The phenotypes of these alleles are annotated with 22 unique terms, many of which group under: embryonic abdomen; embryonic segment; anatomical structure; embryonic tagma; organ system; egg; cell part; abdominal segment 7; blastoderm embryo; spermatozoon; cytoplasmic part; embryonic abdominal segment 4. It has one annotated transcript and one annotated polypeptide.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	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).
Detailed Mapping Data
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)   (Boswell, 1995) 57C7-57C8 (determined by in situ hybridisation)   (Golumbeski et al., 1991)
Experimentally Determined Recombination Data
Location	2-97 (Tearle and Nusslein-Volhard, 1987, Schupbach and Wieschaus, 1989)
Left of (cM)
Right of (cM)
Notes
Molecular Map Data
Gene Order (in direction of increasing cytology) References Gene Order (overall orientation not stated) References Overall orientation not stated: anon-57Cb+ anon-57Cc- anon-57Cd? anon-57Ce? anon-57Cf? anon-57Cg? anon-57Ch? anon-57Ci? anon-57Cj? anon-57Cb+ Pu+ anon-57Cl- Mst57Ca- Pu+ tud+ (O'Donnell et al., 1989)
Gene Model & Products
Please see the GBrowse view of Dmel\tud for information on other features
Comments on Gene Model
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) tud-RA FBtr0071582  NM_057425   8205   2515
Additional Transcript Data & Comments
Reported size (kB)	8.0 (northern blot) (Golumbeski et al., 1991)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kD) Length (aa) Theoretical pI RefSeq ID GenBank protein tud-PA   FBpp0071508   285.3   2515   6.40     NP_476773   AAF46693
Additional Polypeptide Data & Comments
Reported size (kD)	2515 (aa); 285 (kD predicted) (Golumbeski et al., 1991) 2515 (aa); 285 (kD observed) (Bardsley et al., 1993)
Comments
External Data
Linkouts	PANTHER - Protein classification by function, families, and pathways • FBgn0003891
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Tudor (IPR002999) Maternal tudor protein (IPR008191)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA941957   AC008231   AE003453 AAF46693   AI063300   AI135846   AI514276   AI520476   AI541603   BF490958   BI581179   BI581952   BI581980   BI582357   BP546162   BP550857   BZ665735   CO279438   CO280654   CO294509   CO299957   CO300350   CO305501   CO315559   CO328786   CO340674   CZ466740   CZ466741   CZ482422   CZ485298   CZ485299   EC061917   EC062697   EC072210   EC073924   EC075952   EC089265   EC202493   EC212732   EC226865   EC245625   EC247843   EC249260   EC249981   EC258913   EC260356   EC266969   EL871940   X62420 CAA44286   Z50578
UniProtKB/Swiss-Prot	P25823
UniProtKB/TrEMBL
Maps to	FBcl0297751 (FlyBase, 1992-)
Does NOT map to
Identified with	GH27372 (BDGP Project Members, 2000-)
Mapped Features & Mutations
Please see GBrowse or insertion reports for information on insertions of transgenic constructs and features not listed here Type Symbol & Location Additional Notes References protein binding site tud-protein_bind-1 2R:17,070,904..17,070,915 evidence=experimental comment=TC box at -22 to -33 with respect to tud gene transcription start site 2, bound by Trf protein in the presence of TfIIA-L/TfIIA-S or TfIIB proteins bound_moiety=TfIIA-S-XP bound_moiety=TfIIA-L-P1 bound_moiety=Trf[+]P224 (Bergman et al., 2004, Holmes and Tjian, 2000)
External Data
Linkouts	DEDB - Drosophila exon database: splicing graphs • 1180
Crossreferences
Expression Data
FlyBase-Curated Data
Transcript and Protein data	Please see the FlyBase Gene Expression Report for details of gene expression from the literature.
Summary of Transcript Expression
Stage Tissue/Position Reference  oogenesis stage,adult stage | stage S10-S14  follicle cell | posterior (Golumbeski et al., 1991)  oogenesis stage,adult stage  germarium (Golumbeski et al., 1991)  oogenesis stage,adult stage | stage S1-S3  ovariole (Golumbeski et al., 1991)
Marker for
Subcellular Localization
CV Term
Summary of Polypeptide Expression
Stage Tissue/Position Reference  oogenesis stage | stage S7-S9  oocyte (Bardsley et al., 1993)  embryonic stage  mitochondrion (Bardsley et al., 1993)  oogenesis stage,adult stage | stage S10  vitelline membrane (Bardsley et al., 1993)
Marker for
Subcellular Localization
CV Term	nucleus (Bardsley et al., 1993)
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003891 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • FBgn0003891 GEO (NCBI) - Gene expression data: microarray and other high-throughput technologies • FBgn0003891
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal | maternal effect | recessive tud1 tudunspecified lethal | embryonic stage | maternal effect | recessive tudtux46 lethal | embryonic stage | maternal effect | partially | recessive tudunspecified lethal | embryonic stage | germ-line clone | non-rescuable maternal effect | recessive tud2R-26-9 tud2R-415-19 tud2R-78-11 Sterility Allele sterile tudunspecified female sterile | recessive tudunspecified male sterile | maternal effect tud1 tud2 tud3 tud4 male sterile | maternal effect | recessive tud1 male fertile tudtux46 grandchildless tudB42 tudB45 grandchildless (with tudB42) tudB45 grandchildless (with tudB45) tudB42 grandchildless | recessive tud2R-415-19 tudtux46 tudunspecified Phenotype manifest in Allele spermatozoon tud1 pole granule tud1 pole cell tud1 abdominal segment tud1 egg tud1 embryonic abdominal segment | maternal effect (with Df(2R)PurP133a) tud1 polar granule (with Df(2R)PF1) tud1 embryo | germ-line clone | maternal effect | posterior tud2R-26-9 tud2R-415-19 tud2R-78-11 pole granule | maternal effect tudunspecified pole cell | maternal effect tudunspecified tudtux46 embryonic abdominal segment | maternal effect tudunspecified tudC30 tudC73 tudtux46 pole plasm | maternal effect tudunspecified embryonic abdominal segment 1 | maternal effect tudunspecified embryonic abdominal segment 2 | maternal effect tudunspecified embryonic abdominal segment 3 | maternal effect tudunspecified embryonic abdominal segment 4 | maternal effect tudunspecified embryonic abdominal segment 5 | maternal effect tudunspecified embryonic abdominal segment 6 | maternal effect tudunspecified embryonic abdominal segment 7 | maternal effect tudunspecified embryonic abdominal segment 8 | maternal effect tudunspecified embryonic abdominal segment 9 | maternal effect tudunspecified embryonic abdominal segment 10 | maternal effect tudunspecified embryonic abdominal segment 11 | maternal effect tudunspecified germ cell | maternal effect (with Df(2R)PurP133a) tudA7 tudA36 polar granule | maternal effect tudA36 tudB42 tudB45 tudtux46 germ cell | maternal effect tudB42 tudB45 germ cell | maternal effect (with tudB45) tudB42 germ cell | maternal effect (with tudB42) tudB45 gonad | maternal effect tudtux46 polar granule | maternal effect (with Df(2R)PF1) tudtux46
Classical Alleles ( 33 )
For All Classical Alleles Show
Allele of tud Class Mutagen Stocks Known lesion tud1 hypomorph ethyl methanesulfonate 6 Yes tud2 ethyl methanesulfonate X ray 0 -- tud2R-26-9 ethyl methanesulfonate 0 -- tud2R-415-19 ethyl methanesulfonate 0 -- tud2R-78-11 ethyl methanesulfonate 0 -- tud3 ethyl methanesulfonate 0 -- tud4 ethyl methanesulfonate 0 Yes tud5 ethyl methanesulfonate X ray 0 Yes tud6 ethyl methanesulfonate X ray 0 Yes tudA10 ethyl methanesulfonate 0 -- tudA14 ethyl methanesulfonate 0 -- tudA1 ethyl methanesulfonate 0 -- tudA2 0 -- tudA36 ethyl methanesulfonate 0 Yes tudA47 ethyl methanesulfonate 0 -- tudA5 0 -- tudA7 ethyl methanesulfonate 0 Yes tudB42 ethyl methanesulfonate 0 Yes tudB45 ethyl methanesulfonate 0 Yes tudBM-3 0 -- tudC11 ethyl methanesulfonate 0 -- tudC30 ethyl methanesulfonate 0 -- tudC3 ethyl methanesulfonate 0 -- tudC53 ethyl methanesulfonate 0 -- tudC67 ethyl methanesulfonate 0 -- tudC73 ethyl methanesulfonate 0 -- tudC8 ethyl methanesulfonate 0 -- tudKG10175 P-element activity 1 -- tudc00814 piggyBac transposase 1 -- tude04651 piggyBac transposase 1 -- tudf02752 piggyBac transposase 1 -- tudtux46 P-element activity 0 Yes tudunspecified ethyl methanesulfonate 0 --
Alleles Carried on Transgenic Constructs ( 5 )
For All Alleles Carried on Transgenic Constructs Show
Allele of tud Class Mutagen Stocks Known lesion tudGD14219 in vitro construct | RNAi in vitro construct | regulatory fusion 1 Yes tudnos.T:Ivir\HA1 in vitro construct | regulatory fusion in vitro construct | coding region fusion 0 Yes tudΔ1.nos.T:Ivir\HA1 in vitro construct | regulatory fusion in vitro construct | deletion in vitro construct | coding region fusion 0 Yes tudΔ2.nos.T:Ivir\HA1 in vitro construct | regulatory fusion in vitro construct | deletion in vitro construct | coding region fusion 0 Yes tudΔ3.nos.T:Ivir\HA1 in vitro construct | regulatory fusion in vitro construct | deletion in vitro construct | coding region fusion 0 Yes
Aneuploid Aberrations
Useful deficiency	Df(2R)PF1
Useful duplication	Dp(2;2)l-85
Disrupted in	Df(2R)AA21 (O'Donnell et al., 1989) Df(2R)Exel6072 (Ryder, 2004.4.26) Df(2R)Exel6074 (Ryder, 2004.4.26) Df(2R)K11 (O'Donnell et al., 1989) Df(2R)MP1 (O'Donnell et al., 1989) Df(2R)PF1 (O'Donnell et al., 1989) Df(2R)PI12 (O'Donnell et al., 1989) Df(2R)PI13 (Schupbach and Wieschaus, 1989) Df(2R)PK1 (O'Donnell et al., 1989) Df(2R)PL3 (Schupbach and Wieschaus, 1986, O'Donnell et al., 1989) Df(2R)Pu-D17 (Schupbach and Wieschaus, 1986, Schupbach and Wieschaus, 1989, O'Donnell et al., 1989) Df(2R)PurP133a (Tearle and Nusslein-Volhard, 1987, Wang and Lehmann, 1991) In(2R)B46 (Golumbeski et al., 1991) In(2R)tud5 (Golumbeski et al., 1991)
Not disrupted in	Df(2R)CC2 (O'Donnell et al., 1989) Df(2R)F36 (O'Donnell et al., 1989) Df(2R)PC18 (O'Donnell et al., 1989)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD14219} NA characterization construct P{nos-mini-tud.Δ1} NA P{nos-mini-tud.Δ2} NA P{nos-mini-tud.Δ3} NA P{nos-tud.HA} NA
Insertions
	Type of insertions Name Expression data miscellaneous insertions PBac{PB}tudc00814 NA PBac{RB}tude04651 NA PBac{WH}tudf02752 NA P{SUPor-P}tudKG10175 NA insertion of enhancer trap P{lacW}l(2)SH1835SH1835 No
Related Comments
	Please look at the allele reports for the complete phenotype data Mutations in tud cause failure of germ cell formation and deletions in the abdominal segments in the embryo. (Schupbach and Wieschaus, 1986) Mutation in tud results in a maternal effect "grandchildless knirps-like" phenotype. (Schupbach and Wieschaus, 1989) Grandchildless-knirps class of genes. (Hazelrigg et al., 1990) Mutations in maternal posterior class gene tud do not interact with RpII140wimp. (Parkhurst and Ish-Horowicz, 1991) 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. (Chapman et al., 1993) 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. (Bardsley et al., 1993) 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. (Keller, 1995)
Gene Ontology: Function, Process & Cellular Component ( 11 )
Molecular Function
CV term Evidence References
nucleic acid binding inferred from electronic annotation with InterPro:IPR002999 (FlyBase Curators et al., 2004-)
Biological Process
CV term Evidence References
embryonic development traceable author statement (Grau and Gutzeit, 1990) inferred from mutant phenotype (Arkov et al., 2006)
germ cell development traceable author statement (Mahowald, 2001)
intracellular mRNA localization traceable author statement (Wylie, 1999)
mitochondrial rRNA export from mitochondrion inferred from mutant phenotype (Amikura et al., 2001)
polar granule organization and biogenesis inferred from mutant phenotype (Thomson and Lasko, 2004) inferred from mutant phenotype (Arkov et al., 2006)
pole cell development inferred from mutant phenotype (Thomson and Lasko, 2004)
pole cell fate determination non-traceable author statement (Swiss-Prot Project Members, 1992.5.1)
pole plasm assembly inferred from mutant phenotype (Raff et al., 1990) traceable author statement (Breitwieser et al., 1996) traceable author statement (Williamson and Lehmann, 1996) traceable author statement (Mahowald, 2001)
Cellular Component
CV term Evidence References
mitochondrion traceable author statement (Williamson and Lehmann, 1996)
polar granule traceable author statement (Breitwieser et al., 1996) traceable author statement (Williamson and Lehmann, 1996) traceable author statement (Mahowald, 2001) inferred from direct assay (Arkov et al., 2006)