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Gene Dmel\mud

General Information
Symbol	Dmel\mud	Species	D. melanogaster
Name	mushroom body defect	Annotation symbol	CG12047
Feature type	protein_coding_gene	FlyBase ID	FBgn0002873
Gene Model Status	Current	Stock availability	6 publicly available
Genomic Location
Chromosome (arm)	X	Recombination map	1-50
Cytogenetic map	12E5-12E6	Sequence location	X:14,141,876..14,152,628 [+]
Genomic Maps Select View:Gene Models/EvidenceExpression/RegulationMegaViewGene Disruptions, Stocks and Reagents modENCODE 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 mushroom body defect is referred to in FlyBase by the symbol Dmel\mud (CG12047, FBgn0002873). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: protein binding; DNA binding. There is experimental evidence that it is involved in the biological process: asymmetric neuroblast division; spindle assembly involved in female meiosis II; mushroom body development. 13 alleles are reported. The phenotypes of these alleles are annotated with: somatic stem cell; organ system subdivision; larval abdominal segment; multi-cell-component structure; adult segment; abdominal segment 7; somatic cell; nervous system; larval abdominal segment 5; abdominal 2 neuroblast ventro-medial cluster; sensillum; abdominal 5 neuroblast ventro-medial cluster; FBbt:00003689; organelle. It has 5 annotated transcripts and 5 annotated polypeptides. Protein features are: Manganese/iron superoxide dismutase; Nuclear hormone receptor, ligand-binding; Prefoldin; RuvA domain 2-like. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of high expression to a trough of very low expression. Peak expression observed within 00-06 hour embryonic stages. Summary of FlyAtlas Anatomical Expression Data: Two or more Affy2 ProbeSets identify exons of this gene. This is a summary of the tissue expression peaks exhibited in at least one of these ProbeSets. Expression at moderate levels in the following post-embryonic organs or tissues: adult ovary, adult testis. Gene sequence location is X:14141876..14152628.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	mud: mushroom body defect (J.C. Hall) Neuropile of mushroom bodies (usually in dorsal brain) is missing (Heisenberg, 1980); penetrance for this anatomical phenotype (on which criterion the mutant was isolated) ca. 90%. Calyces that should innervate pedunculi and lobes of mushroom bodies are enlarged, as are antennal lobes in anterior brain. At sites of calyces, large numbers of thin axons form distinct lobes outside main neuropile of brain. Viability of mutant females poor, and they are sterile as well. Development of mushroom bodies apparently normal until pupation, when "reconstruction" of this bilaterally paired brain entity (this process being a normal feature of wild-type metamorphosis) is aberrant (Technau and Heisenberg, 1982). Supernumerary neuroblasts observed in the larval brain, owing to anomalous proliferation of such cells postembryonically.
Recent Updates
Description	What does this section display? This section contains items that were added to this record for each release. It currently only tracks new links between this FlyBase report and other FlyBase data classes (e.g. genes, references, stocks) or controlled vocabulary terms (e.g. GO, anatomy terms). What does this section not display? This section does not currently display links that were removed or gene model changes.
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FB2012_01	References Wee et al., 2011, J. Cell Biol. 195(3): 369--376 Cabernard et al., 2010, Nature 467(7311): 91--94 Capalbo et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(42): 17343--17348 Sequence features BKN23701
FB2011_10
All updates	Click here to see a list of all updates to this record from FB2010_08 and on.
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 12E5-12E6   Limits computationally determined from genome sequence between P{EP}l(1)G0007EP764 and P{EP}EP1599
Experimentally Determined Cytological Location
Cytogenetic map Notes References 12E-12E     (Guan, 1999.7.31)
Experimentally Determined Recombination Data
Location	1-50 (Heisenberg, 1980)
Left of (cM)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\mud for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
	Gene model reviewed during 5.39 DGC clone LD29039 appears problematic: incomplete CDS Apparent introns within longest coding exon in cDNAs and RNA-seq junctions within the same region are likely to be artifacts; possibly caused by reverse transcriptase template-switching.
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) mud-RF FBtr0308325     7827   2394 mud-RG FBtr0308326     6177   1933 mud-RH FBtr0308327     8346   2567 mud-RI FBtr0308328     8532   2113 mud-RJ FBtr0308329     6986   2165
Additional Transcript Data & Comments
Reported size (kB)	6.003 (longest cDNA) (Guan et al., 2000)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein mud-PF   FBpp0300644   274.5   2394   5.47       mud-PG   FBpp0300645   223.3   1933   5.02       mud-PH   FBpp0300646   294.1   2567   5.46       mud-PI   FBpp0300647   242.8   2113   5.06       mud-PJ   FBpp0300648   248.9   2165   5.14
Additional Polypeptide Data & Comments
Reported size (kDa)	2501, 1871 (aa) (Guan et al., 2000)
Comments	Each mud protein isoform is predicted to have a long coiled-coil in the central region, and a carboxy-terminal transmembrane domain. The region predicted to form a coiled-coil includes a segment of 42 amino acids that is repeated 10 times. The 1871 amino acid isoform is predicted to have a single transmembrane segment, whereas the two longer isoforms are predicted to have two transmembrane segments each. (Guan et al., 2000, Guan et al., 2000, Guan et al., 2000)
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Manganese/iron superoxide dismutase (IPR001189) Prefoldin (IPR009053) RuvA domain 2-like (IPR010994) Nuclear hormone receptor, ligand-binding (IPR008946)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA392172   AA820568   AA949597   AA951447   AA952024   AF174134 AAF88146   AF209068 AAF89163   AI455461   AI512725   AW942926   AY058595 AAL13824   BF496125   BF501328   BF503763   BI228050   BI230863   BI355966   BP542150   BT024978   CO179303   CO262133   CO266443   CO270264   CO273644   CO279851   CO291228   CO311485   CO315085   CO328916   CO333093   CZ483683   DQ465527 ABF00987   EC065548   EC070875   EC070876   EC073022   EC207312   EC217166   EC219387   EC221174   EC232491   EC242466   EC251218   EL874863   EL874864   EL874992   EL874993   EV576929   EV579655
UniProtKB/Swiss-Prot
UniProtKB/TrEMBL	Q1KN24 Q8IR54 Q8IR55 Q95TR6 Q9NCL3 Q9NCW7 Q9VY43
Mapped Features
Mapped Features have been reorganized, please see this article for details. Additional mapped features and mutations can be found on GBrowse or related reports. Type Symbol & Location Additional Notes References
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage     (Guan et al., 2000) larval stage     (Guan et al., 2000) larval stage | male     (Guan et al., 2000) adult stage     (Guan et al., 2000)
Additional Descriptive Data
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
mass spectroscopy Stage Tissue/Position (including subcellular localization) Reference day 5 of adulthood & male -- day 7 of adulthood & male   spermatozoon   (Wasbrough et al., 2010)
Additional Descriptive Data
Marker for
Subcellular Localization
CV Term
Notes
High-Throughput Expression Data
or
See Gelbart and Emmert, 2010.10.13 for analysis details and data files for all genes. modENCODE Temporal Expression Data for FBgn0002873    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0002873 Very low expression bin max Moderate expression bin max High expression bin max Extremely high expression bin max Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of high expression to a trough of very low expression. Peak expression observed within 00-06 hour embryonic stages. [download data (TSV)] Guide to modENCODE expression level colors   No expression (0 - 0)   Extremely low expression (1 - 10)   Very low expression (11 - 100)   Low expression (101 - 400)   Moderate expression (401 - 1400)   Moderately high expression (1401 - 4000)   High expression (4001 - 10000)   Very high expression (10001 - 100000)   Extremely high expression (100001 - 2000000) Linear, scaled to maximum FBgn0002873 expression level Developmental Stage   Expression Level embryo 00-02hr    3433 embryo 02-04hr    4517 embryo 04-06hr    2582 embryo 06-08hr    1917 embryo 08-10hr    1264 embryo 10-12hr    246 embryo 12-14hr    490 embryo 14-16hr    314 embryo 16-18hr    218 embryo 18-20hr    116 embryo 20-22hr    378 embryo 22-24hr    371 larva L1    48 larva L2    59 larva L3 12hr old    115 larva L3 puffstage 1-2    312 larva L3 puffstage 3-6    713 larva L3 puffstage 7-9    895 white prepupae new    769 white prepupae 12hr    1469 white prepupae 24hr    1502 pupae 2d postWPP    1079 pupae 3d postWPP    800 pupae 4d postWPP    551 adult male 01day    1088 adult male 05day    1700 adult male 30day    1128 adult female 01day    1260 adult female 05day    2650 adult female 30day    2498 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (3433) embryo 02-04hr  (4517) embryo 04-06hr  (2582) embryo 06-08hr  (1917) embryo 08-10hr  (1264) embryo 10-12hr  (246) embryo 12-14hr  (490) embryo 14-16hr  (314) embryo 16-18hr  (218) embryo 18-20hr  (116) embryo 20-22hr  (378) embryo 22-24hr  (371) larva L1    48 larva L2    59 larva L3 12hr old  (115) larva L3 puffstage 1-2  (312) larva L3 puffstage 3-6  (713) larva L3 puffstage 7-9  (895) white prepupae new  (769) white prepupae 12hr  (1469) white prepupae 24hr  (1502) pupae 2d postWPP  (1079) pupae 3d postWPP  (800) pupae 4d postWPP  (551) adult male 01day  (1088) adult male 05day  (1700) adult male 30day  (1128) adult female 01day  (1260) adult female 05day  (2650) adult female 30day  (2498) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (3433) embryo 02-04hr  (4517) embryo 04-06hr  (2582) embryo 06-08hr  (1917) embryo 08-10hr    1264 embryo 10-12hr    246 embryo 12-14hr    490 embryo 14-16hr    314 embryo 16-18hr    218 embryo 18-20hr    116 embryo 20-22hr    378 embryo 22-24hr    371 larva L1    48 larva L2    59 larva L3 12hr old    115 larva L3 puffstage 1-2    312 larva L3 puffstage 3-6    713 larva L3 puffstage 7-9    895 white prepupae new    769 white prepupae 12hr  1469 white prepupae 24hr  1502 pupae 2d postWPP    1079 pupae 3d postWPP    800 pupae 4d postWPP    551 adult male 01day    1088 adult male 05day  (1700) adult male 30day    1128 adult female 01day    1260 adult female 05day  (2650) adult female 30day  (2498) Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    3433 embryo 02-04hr    4517 embryo 04-06hr    2582 embryo 06-08hr    1917 embryo 08-10hr    1264 embryo 10-12hr    246 embryo 12-14hr    490 embryo 14-16hr    314 embryo 16-18hr    218 embryo 18-20hr    116 embryo 20-22hr    378 embryo 22-24hr    371 larva L1    48 larva L2    59 larva L3 12hr old    115 larva L3 puffstage 1-2    312 larva L3 puffstage 3-6    713 larva L3 puffstage 7-9    895 white prepupae new    769 white prepupae 12hr    1469 white prepupae 24hr    1502 pupae 2d postWPP    1079 pupae 3d postWPP    800 pupae 4d postWPP    551 adult male 01day    1088 adult male 05day    1700 adult male 30day    1128 adult female 01day    1260 adult female 05day    2650 adult female 30day    2498 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    3433 embryo 02-04hr    4517 embryo 04-06hr    2582 embryo 06-08hr    1917 embryo 08-10hr    1264 embryo 10-12hr    246 embryo 12-14hr    490 embryo 14-16hr    314 embryo 16-18hr    218 embryo 18-20hr    116 embryo 20-22hr    378 embryo 22-24hr    371 larva L1    48 larva L2    59 larva L3 12hr old    115 larva L3 puffstage 1-2    312 larva L3 puffstage 3-6    713 larva L3 puffstage 7-9    895 white prepupae new    769 white prepupae 12hr    1469 white prepupae 24hr    1502 pupae 2d postWPP    1079 pupae 3d postWPP    800 pupae 4d postWPP    551 adult male 01day    1088 adult male 05day    1700 adult male 30day    1128 adult female 01day    1260 adult female 05day    2650 adult female 30day    2498 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0002873 expression level Developmental Stage   Expression Level embryo 00-02hr    3433 embryo 02-04hr    4517 embryo 04-06hr    2582 embryo 06-08hr    1917 embryo 08-10hr    1264 embryo 10-12hr    246 embryo 12-14hr    490 embryo 14-16hr    314 embryo 16-18hr    218 embryo 18-20hr    116 embryo 20-22hr    378 embryo 22-24hr    371 larva L1    48 larva L2    59 larva L3 12hr old    115 larva L3 puffstage 1-2    312 larva L3 puffstage 3-6    713 larva L3 puffstage 7-9    895 white prepupae new    769 white prepupae 12hr    1469 white prepupae 24hr    1502 pupae 2d postWPP    1079 pupae 3d postWPP    800 pupae 4d postWPP    551 adult male 01day    1088 adult male 05day    1700 adult male 30day    1128 adult female 01day    1260 adult female 05day    2650 adult female 30day    2498 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (3433) embryo 02-04hr  (4517) embryo 04-06hr  (2582) embryo 06-08hr  (1917) embryo 08-10hr  (1264) embryo 10-12hr  (246) embryo 12-14hr  (490) embryo 14-16hr  (314) embryo 16-18hr  (218) embryo 18-20hr  116 embryo 20-22hr  (378) embryo 22-24hr  (371) larva L1    48 larva L2    59 larva L3 12hr old  115 larva L3 puffstage 1-2  (312) larva L3 puffstage 3-6  (713) larva L3 puffstage 7-9  (895) white prepupae new  (769) white prepupae 12hr  (1469) white prepupae 24hr  (1502) pupae 2d postWPP  (1079) pupae 3d postWPP  (800) pupae 4d postWPP  (551) adult male 01day  (1088) adult male 05day  (1700) adult male 30day  (1128) adult female 01day  (1260) adult female 05day  (2650) adult female 30day  (2498) Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (3433) embryo 02-04hr  (4517) embryo 04-06hr  (2582) embryo 06-08hr  1917 embryo 08-10hr    1264 embryo 10-12hr    246 embryo 12-14hr    490 embryo 14-16hr    314 embryo 16-18hr    218 embryo 18-20hr    116 embryo 20-22hr    378 embryo 22-24hr    371 larva L1    48 larva L2    59 larva L3 12hr old    115 larva L3 puffstage 1-2    312 larva L3 puffstage 3-6    713 larva L3 puffstage 7-9    895 white prepupae new    769 white prepupae 12hr  1469 white prepupae 24hr  1502 pupae 2d postWPP    1079 pupae 3d postWPP    800 pupae 4d postWPP    551 adult male 01day    1088 adult male 05day  1700 adult male 30day    1128 adult female 01day    1260 adult female 05day  (2650) adult female 30day  (2498) Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    3433 embryo 02-04hr    4517 embryo 04-06hr    2582 embryo 06-08hr    1917 embryo 08-10hr    1264 embryo 10-12hr    246 embryo 12-14hr    490 embryo 14-16hr    314 embryo 16-18hr    218 embryo 18-20hr    116 embryo 20-22hr    378 embryo 22-24hr    371 larva L1    48 larva L2    59 larva L3 12hr old    115 larva L3 puffstage 1-2    312 larva L3 puffstage 3-6    713 larva L3 puffstage 7-9    895 white prepupae new    769 white prepupae 12hr    1469 white prepupae 24hr    1502 pupae 2d postWPP    1079 pupae 3d postWPP    800 pupae 4d postWPP    551 adult male 01day    1088 adult male 05day    1700 adult male 30day    1128 adult female 01day    1260 adult female 05day    2650 adult female 30day    2498 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    3433 embryo 02-04hr    4517 embryo 04-06hr    2582 embryo 06-08hr    1917 embryo 08-10hr    1264 embryo 10-12hr    246 embryo 12-14hr    490 embryo 14-16hr    314 embryo 16-18hr    218 embryo 18-20hr    116 embryo 20-22hr    378 embryo 22-24hr    371 larva L1    48 larva L2    59 larva L3 12hr old    115 larva L3 puffstage 1-2    312 larva L3 puffstage 3-6    713 larva L3 puffstage 7-9    895 white prepupae new    769 white prepupae 12hr    1469 white prepupae 24hr    1502 pupae 2d postWPP    1079 pupae 3d postWPP    800 pupae 4d postWPP    551 adult male 01day    1088 adult male 05day    1700 adult male 30day    1128 adult female 01day    1260 adult female 05day    2650 adult female 30day    2498 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Developmental Stage   Expression Level embryo 00-02hr     embryo 02-04hr     embryo 04-06hr     embryo 06-08hr     embryo 08-10hr     embryo 10-12hr     embryo 12-14hr     embryo 14-16hr     embryo 16-18hr     embryo 18-20hr     embryo 20-22hr     embryo 22-24hr     larva L1     larva L2     larva L3 12hr old     larva L3 puffstage 1-2     larva L3 puffstage 3-6     larva L3 puffstage 7-9     white prepupae new     white prepupae 12hr     white prepupae 24hr     pupae 2d postWPP     pupae 3d postWPP     pupae 4d postWPP     adult male 01day     adult male 05day     adult male 30day     adult female 01day     adult female 05day     adult female 30day     FlyAtlas Anatomical Expression Data for FBgn0002873    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0002873 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Two or more Affy2 ProbeSets identify exons of this gene. This is a summary of the tissue expression peaks exhibited in at least one of these ProbeSets. Expression at moderate levels in the following post-embryonic organs or tissues: adult ovary, adult testis. [download data (TSV)] Guide to FlyAtlas expression level colors   No expression (0 - 9.999)   Low expression (10 - 99.999)   Moderate expression (100 - 499.999)   High level expression (500 - 999.999)   Very high expression (1000 - 25000) Heatmap Tissue   Expression Level Larval Central Nervous System no informative data Larval Midgut     Larval Hindgut     Larval Malpighian Tubules     Larval Fat Body no informative data Larval Salivary Gland     Larval Trachea     Larval Carcass     Adult Head     Adult Eye     Adult Brain     Adult Thoracic-Abdominal Ganglion     Adult Crop     Adult Midgut     Adult Hindgut     Adult Malpighian Tubules     Adult Fat Body     Adult Salivary Gland     Adult Heart     Adult VirginFemale Spermatheca     Adult InseminatedFemale Spermatheca     Adult Ovary no informative data Adult Testis no informative data Adult Male Accessory Gland     Adult Carcass     modENCODE Temporal Expression Data (Graveley et al., 2011) FlyAtlas Anatomical Expression Data (Chintapalli et al., 2007)
Expression Clusters
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0002873 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG12047-RC;CG12047-RC;CG12047-RA
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal | embryonic stage | non-rescuable maternal effect | recessive mud4 lethal | female mud1 mud3 semi-lethal mud1 mud2 mud3 mud4 semi-lethal | female limited mud1 mud4 Sterility Allele female sterile mud1 mud2 mud3 mud4 male fertile mud1 mud2 mud3 mud4 sterile (with Df(1)KA9) mud1 Other Phenotypes Allele chemical sensitive mud4 chemical sensitive (with mud1) mud4 chemical sensitive (with mud4) mud1 chemical sensitive | codominant mud4 locomotor behavior defective mud1 mud4 wild-type mud+t18 mud+tcos24 Phenotype manifest in Allele abdominal 2 neuroblast dorso-lateral cluster mud2 abdominal 2 neuroblast ventro-medial cluster mud2 abdominal 3 neuroblast dorso-lateral cluster mud2 abdominal 3 neuroblast ventro-medial cluster mud2 abdominal 4 neuroblast dorso-lateral cluster mud2 abdominal 4 neuroblast ventro-medial cluster mud2 abdominal 5 neuroblast dorso-lateral cluster mud2 abdominal 5 neuroblast ventro-medial cluster mud2 abdominal 6 neuroblast dorso-lateral cluster mud2 abdominal 6 neuroblast ventro-medial cluster mud2 abdominal 7 neuroblast dorso-lateral cluster mud2 abdominal 7 neuroblast ventro-medial cluster mud2 adult central complex mud1 mud4 adult mushroom body mud4 antennal lobe mud1 egg & chromatin | maternal effect mud4 egg & spindle | maternal effect mud4 egg | maternal effect mud4 ganglion mother cell | somatic clone (with mud3) mud4 ganglion mother cell | somatic clone (with mud4) mud3 Kenyon cell mud1 mud4 lobe system of mushroom body mud1 mud4 macrochaeta mudBG02499 mushroom body mud1 mud2 mud4 mushroom body calyx mud1 mud2 mud4 mushroom body medial lobe mud4 mushroom body pedunculus mud2 neuroblast | somatic clone | supernumerary (with mud3) mud4 neuroblast | somatic clone | supernumerary (with mud4) mud3 neuroblast | supernumerary (with mud3) mud4 neuroblast | supernumerary (with mud4) mud3 neuropil mud1 spindle (with mud3) mud4 spindle (with mud4) mud3 vertical branch of mushroom body mud4
Classical Alleles ( 7 )
For All Classical Alleles Show
Allele of mud Class Mutagen Stocks Known lesion mud1 ethyl methanesulfonate 2 Yes mud4 ethyl methanesulfonate 1 -- mudf01205 piggyBac transposase 1 -- mud2 PM hybrid dysgenesis 0 Yes mud3 ethyl methanesulfonate 0 Yes mudBG02499 P-element activity 0 Yes mudunspecified 0 --
Alleles Carried on Transgenic Constructs ( 6 )
For All Alleles Carried on Transgenic Constructs Show
Allele of mud Class Mutagen Stocks Known lesion mudHMS01458 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes mudJF02911 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes mud+t18 in vitro construct - genomic fragment 0 Yes mud+tcos24 in vitro construct - genomic fragment 0 Yes mudGD3353 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes mudpJMD1 in vitro construct - genomic fragment 0 Yes
Aneuploid Aberrations
Disrupted in	Df(1)benCO1 (Guan et al., 2000) Df(1)benCO2 (Guan et al., 2000) Df(1)BSC628 (Christensen et al., 2008.9.29) Df(1)BSC705 (Christensen et al., 2008.12.28) Df(1)BSC857 (Cook et al., 2009.11.25) Df(1)KA9 (Guan et al., 2000) Df(1)RK3 (Guan et al., 2000)
Not disrupted in	Df(1)g (Guan et al., 2000) Df(1)NP5 (Guan et al., 2000, Mitchell, 1997.2.17) Df(1)RK5 (Guan et al., 2000) T(1;Y)B146 (Scott, 2008.5.20)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD3353} NA P{TRiP.HMS01458} NA P{TRiP.JF02911} NA characterization construct P{mud.cos24} NA P{mud.pJMD1} NA P{mud.pZHRC} NA
Insertions
	Type of insertions Name Expression data miscellaneous insertions P{GT1}mudBG02499 NA PBac{WH}mudf01205 NA insertion of mobile activating element P{GSV1}EP-689EP-689 NA
Gene Ontology: Function, Process & Cellular Component ( 19 unique terms )
Terms Based on Experimental Evidence ( 10 terms )
Molecular Function
CV term Evidence References
DNA binding inferred from direct assay (Smith and Prehoda, 2011) protein binding inferred from physical interaction with raps (Nipper et al., 2007)
Biological Process
CV term Evidence References
asymmetric neuroblast division inferred from mutant phenotype (Cabernard and Doe, 2009) mushroom body development inferred from mutant phenotype (de Belle and Heisenberg, 1996) spindle assembly involved in female meiosis II inferred from mutant phenotype (Yu et al., 2006)
Cellular Component
CV term Evidence References
apical cortex inferred from direct assay (Nipper et al., 2007) basal cortex inferred from direct assay (Nipper et al., 2007) centrosome inferred from direct assay (Nipper et al., 2007) microtubule associated complex inferred from direct assay (Hughes et al., 2008) spindle midzone inferred from direct assay (Yu et al., 2006)
Terms Based on Predictions or Assertions ( 10 terms )
Molecular Function
CV term Evidence References
metal ion binding inferred from electronic annotation with InterPro:IPR001189 (FlyBase Curators et al., 2004-) sequence-specific DNA binding transcription factor activity inferred from electronic annotation with InterPro:IPR008946 (FlyBase Curators et al., 2004-) steroid hormone receptor activity inferred from electronic annotation with InterPro:IPR008946 (FlyBase Curators et al., 2004-) superoxide dismutase activity inferred from electronic annotation with InterPro:IPR001189 (FlyBase Curators et al., 2004-)
Biological Process
CV term Evidence References
mushroom body development traceable author statement (Guan et al., 2000) oxidation-reduction process inferred from electronic annotation with InterPro:IPR001189 (FlyBase Curators et al., 2004-) regulation of transcription, DNA-dependent inferred from electronic annotation with InterPro:IPR008946 (FlyBase Curators et al., 2004-) steroid hormone mediated signaling pathway inferred from electronic annotation with InterPro:IPR008946 (FlyBase Curators et al., 2004-) superoxide metabolic process inferred from electronic annotation with InterPro:IPR001189 (FlyBase Curators et al., 2004-)
Cellular Component
CV term Evidence References
nucleus inferred from electronic annotation with InterPro:IPR008946 (FlyBase Curators et al., 2004-)
Sequence Ontology: Class of Gene
	nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
Protein-protein
Interacting group Assay References
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions mud allele Gene References mud4 w (Campbell and Nash, 2001)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 69318 DroID - A comprehensive database of gene and protein interactions. • FBgn0002873 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 44839
Orthologs
Genome-wide drosophilid orthologs	Dana\GF22554 Dere\GG17839 Dgri\GH12928 Dper\GL14926 Dsim\GD17165 Dvir\GJ19063 Dwil\GK20155
Curated drosophilid orthologs	Dpse\GA11358
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL014706-PA Aedes aegypti (mosquito) AGAP000323-PA Anopheles gambiae (mosquito) XP_001952645.1 Acyrthosiphon pisum (pea aphid) BGIBMGA004128-PA Bombyx mori (silk worm) CPIJ017450 Culex pipens (mosquito) ENSP00000369129 Homo sapiens (human) ENSPTRP00000030221 Pan troglodytes (chimpanzee) ENSRNOP00000028440 Rattus norvegicus (norway rat) TA20220 Theileria annulata (apicomplexan parasite) XP_975536.2 Tribolium castaneum (red flour beetle) 91913.m00053 Trichomonas vaginalis (parasitic flagellated protozoan) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • FBgn0234614 FBgn0234612 FBgn0232609 FBgn0232322 FBgn0232321 FBgn0222152 FBgn0210809 FBgn0206208 FBgn0172576 FBgn0172575 JGI_V11_338968 JGI_V11_335548 JGI_V11_254638 FBgn0247966 FBgn0247961 FBgn0152530 FBgn0138187 FBgn0138186 FBgn0120406 FBgn0110060 FBgn0099548 CPIJ017450 CPIJ017448 CPIJ005801 BGIBMGA004128 ACYPI33638 ACYPI21409 ACYPI002164 AGAP000323 AAEL015340 AAEL014706 AAEL002942 PHUM308510 PHUM353010 GLEAN_10007
Stocks & Reagents
Stocks Listed in FlyBase ( 6 )
Bloomington	35830 mud1/FM7a 9562 mud1/FM7i, P{ActGFP}JMR3 9563 mud4/FM7i, P{ActGFP}JMR3 28074 y1 v1; P{TRiP.JF02911}attP2 35044 y1 sc* v1; P{TRiP.HMS01458}attP2
Harvard	- PBac{WH}mudf01205
Genomic Clones ( 1 )
	BACR12J05 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 38 )
	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	IP14842 LD29039
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	AT04961 AT09842 AT16617 LD11076 LD29552 LD31911 LD44694 RE16121 RE25468 RE37037
Other clones	11081 28724 35305 119419 281012 CK01.1C.D5 CK01.75B.C7 CK01.99D.D12 CK01.108D.E3 Dmel_dig_013101_0649_2794 Dmel_dig_013326_0609_2053 EK059012 EK076748 EK084448 EK098118 EK111723 EK136160 EK146433 EK224707 EK238963 EK291648 EK305651 FGC009F03 IP14942 IP15042 LD24364
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0002873 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 44839
Antibody Information
	polyclonal (Izumi et al., 2006)
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of
Additional comments	Annotation CG12047 split into CG12047 (which corresponds to mud) and CG32599 in release 3 of the genome annotation. (FlyBase Genome Annotators, 2002-2003)
Other Comments
	Expression is enriched in embryonic gonads. (Shigenobu et al., 2006) Mutation of mud interferes with the regulation of central nervous system development by affecting the proliferation pattern. (Prokop and Technau, 1994) Neuropil of mushroom bodies (usually in dorsal brain) is missing (Heisenberg, 1980); penetrance for this anatomical phenotype (on which criterion the mutant was isolated) ca. 90%. Calyces that should innervate pedunculi and lobes of mushroom bodies are enlarged, as are antennal lobes in anterior brain. At sites of calyces, large numbers of thin axons form distinct lobes outside main neuropil of brain. Viability of mutant females poor and they are sterile as well. Development of mushroom bodies apparently normal until pupation, when 'reconstruction' of this bilaterally paired brain entity (this process being a normal feature of wild-type metamorphosis) is aberrant (Technau and Heisenberg, 1982). Supernumerary neuroblasts observed in the larval brain, owing to anomalous proliferation of such cells postembryonically.
External Crossreferences & Linkouts
Sequence Crossreferences
	DDBJ / EMBL / GenBank • AA392172 AA820568 AA949597 AA951447 AA952024 AAF48362 AAF48362.2 AAF88146 AAF89163 AAL13824 AAN09583 AAN09583.1 AAN09583.2 AAN09584 AAN09584.1 ABF00987 AF174134 AF209068 AI455461 AI512725 AW942926 AY058595 CZ483683 DQ465527 Entrez Gene - A searchable database of RefSeq genes. • 44839 UniProtKB/TrEMBL • Q1KN24 Q8IR54 Q8IR55 Q9NCL3 Q9NCW7 Q9VY43 Q95TR6
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Manganese/iron superoxide dismutase (IPR001189) Prefoldin (IPR009053) RuvA domain 2-like (IPR010994) Nuclear hormone receptor, ligand-binding (IPR008946)
Linkouts
	BioGRID - A database of protein and genetic interactions • 69318 DroID - A comprehensive database of gene and protein interactions. • FBgn0002873 DRSC - Results from RNAi screens. • FBgn0002873 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0002873 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG12047-RC;CG12047-RC;CG12047-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0002873 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 44839 InParanoid A subset of ortholog calls from InParanoid. • AAEL014706-PA Aedes aegypti (mosquito) AGAP000323-PA Anopheles gambiae (mosquito) XP_001952645.1 Acyrthosiphon pisum (pea aphid) BGIBMGA004128-PA Bombyx mori (silk worm) CPIJ017450 Culex pipens (mosquito) ENSP00000369129 Homo sapiens (human) ENSPTRP00000030221 Pan troglodytes (chimpanzee) ENSRNOP00000028440 Rattus norvegicus (norway rat) TA20220 Theileria annulata (apicomplexan parasite) XP_975536.2 Tribolium castaneum (red flour beetle) 91913.m00053 Trichomonas vaginalis (parasitic flagellated protozoan) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /hjmuller/mud1.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 44839 modMine - Data generated by the modENCODE project. • FBgn0002873 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • FBgn0234614 FBgn0234612 FBgn0232609 FBgn0232322 FBgn0232321 FBgn0222152 FBgn0210809 FBgn0206208 FBgn0172576 FBgn0172575 JGI_V11_338968 JGI_V11_335548 JGI_V11_254638 FBgn0247966 FBgn0247961 FBgn0152530 FBgn0138187 FBgn0138186 FBgn0120406 FBgn0110060 FBgn0099548 CPIJ017450 CPIJ017448 CPIJ005801 BGIBMGA004128 ACYPI33638 ACYPI21409 ACYPI002164 AGAP000323 AAEL015340 AAEL014706 AAEL002942 PHUM308510 PHUM353010 GLEAN_10007
Synonyms & Secondary IDs ( 10 )
Reported As
Symbol Synonym	CG12047 (Nash, 2000.8.14, Norga et al., 2003, Tzolovsky et al., 2002, Giot et al., 2003, Molnar et al., 2006, Brandt, 2006, Perkins et al., 2009, Hughes et al., 2008, Ni et al., 2011.7.19) KS63   Mud (Izumi et al., 2005, Wang et al., 2006, Lee et al., 2006, Bonaccorsi et al., 2007, Chia et al., 2008, Yamashita and Fuller, 2008, Basto et al., 2008, Hughes et al., 2008, Ségalen et al., 2010, Smith and Prehoda, 2011) mud (Yu et al., 2006, Shigenobu et al., 2006, Cabernard et al., 2007, Helfrich-Forster et al., 2002, Scott, 2008.5.20, Molnar et al., 2006, Gan et al., 2010, Christensen et al., 2008.9.29, Christensen et al., 2008.12.28, Nipper et al., 2007, Bowman et al., 2006, Siller et al., 2006, Izumi et al., 2006, Speicher et al., 2008, Hong et al., 2008, Serway et al., 2009, Cabernard and Doe, 2009, Brandt, 2006, Cook et al., 2009.11.25, McIntyre et al., 2006, Godin et al., 2010, Wasbrough et al., 2010, Cabernard et al., 2010) NuMA (Bowman et al., 2006)
Name Synonym	mushroom bodies deranged (Serway et al., 2009) mushroom body defect (Helfrich-Forster et al., 2002) Mushroom body defect (Speicher et al., 2007, Bonaccorsi et al., 2007, Izumi et al., 2006, Speicher et al., 2008) Mushroom Body Defect (Bowman et al., 2006) mushroom body defective (Chia et al., 2008, Cabernard and Doe, 2009)
Secondary FlyBase IDs
	FBgn0030557 FBgn0052599
References ( 77 )
Generate a list of	All references relating to this gene ( 77 )
List References by type	Research paper  ( 36 ) Supplementary material  ( 4 ) Review  ( 6 ) Personal communication to FlyBase  ( 10 ) Abstract  ( 12 ) FlyBase analysis  ( 2 ) DNA/RNA sequence record  ( 1 ) Computer file  ( 3 ) Automatic genome annotation  ( 1 ) Book  ( 1 ) Letter  ( 1 )
Recent research papers ( 7 )
	Capalbo et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(42): 17343--17348 Rab5 GTPase controls chromosome alignment through Lamin disassembly and relocation of the NuMA-like protein Mud to the poles during mitosis. [FBrf0216458] Smith and Prehoda, 2011, Mol. Cell 43(4): 540--549 Robust spindle alignment in Drosophila neuroblasts by ultrasensitive activation of pins. [FBrf0214708] Wee et al., 2011, J. Cell Biol. 195(3): 369--376 Canoe binds RanGTP to promote PinsTPR/Mud-mediated spindle orientation. [FBrf0216492] Cabernard et al., 2010, Nature 467(7311): 91--94 A spindle-independent cleavage furrow positioning pathway. [FBrf0211664] Gan et al., 2010, Cell Res. 20(7): 763--783 Dynamic regulation of alternative splicing and chromatin structure in Drosophila gonads revealed by RNA-seq. [FBrf0211191] Godin et al., 2010, Neuron 67(3): 392--406 Huntingtin Is Required for Mitotic Spindle Orientation and Mammalian Neurogenesis. [FBrf0211450] Ségalen et al., 2010, Dev. Cell 19(5): 740--752 The Fz-Dsh Planar Cell Polarity Pathway Induces Oriented Cell Division via Mud/NuMA in Drosophila and Zebrafish. [FBrf0212266] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...