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Gene Dmel\Mad

General Information
Symbol	Dmel\Mad	Species	D. melanogaster
Name	Mothers against dpp	Annotation symbol	CG12399
Feature type	protein_coding_gene	FlyBase ID	FBgn0011648
Gene Model Status	Current	Stock availability	14 publicly available
Also Known As	pMad, p-Mad, dMAD
Genomic Location
Chromosome (arm)	2L	Recombination map	2-7.7
Cytogenetic map	23D3-23D3	Sequence location	2L:3,146,056..3,159,643 [-]
Genomic Maps Select View:Gene Models/EvidenceExpression/RegulationMegaViewGene Disruptions, Stocks and Reagents Beta	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene Mothers against dpp is referred to in FlyBase by the symbol Dmel\Mad (CG12399, FBgn0011648). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: protein binding; RNA polymerase II distal enhancer sequence-specific DNA binding; sequence-specific DNA binding; RNA polymerase II distal enhancer sequence-specific DNA binding transcription factor activity; RNA polymerase II activating transcription factor binding; RNA polymerase II transcription coactivator activity. There is experimental evidence for 18 unique biological process terms, many of which group under: biological regulation; single-organism developmental process; post-embryonic organ morphogenesis; regulation of developmental process; multicellular organism reproduction; sensory organ development; stem cell differentiation; appendage development; ovarian follicle cell development; cellular response to endogenous stimulus; positive regulation of cellular component biogenesis; regulation of S phase of mitotic cell cycle; localization; positive regulation of cellular biosynthetic process; trunk segmentation; negative regulation of metabolic process. 91 alleles are reported. The phenotypes of these alleles are annotated with: organ system; adult segment; organ system subdivision; thoracic segment; portion of tissue; imaginal precursor; synapse; tracheal system; embryonic/larval hemocoel; embryonic/larval fat body; epithelial furrow; external compound sense organ. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: Dwarfin; MAD homology 1, Dwarfin-type; MAD homology, MH1; SMAD domain, Dwarfin-type; SMAD domain-like; SMAD/FHA domain. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of high expression to a trough of moderate expression. Peak expression observed at stages throughout embryogenesis. Summary of FlyAtlas Anatomical Expression Data: Nearly all larval and adult tissues/organs expressed at moderate levels. Expression at high levels in the following post-embryonic organs or tissues: adult ovary. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, larval/adult central nervous system, adult crop, larval/adult midgut, larval/adult hindgut, larval/adult Malpighian tubules, adult heart, larval/adult fat body, larval/adult salivary gland, larval trachea, adult spermathecae, adult male accessory gland, larval/adult carcass. Comments on Affy2 ProbeSet: ProbeSet 1634683_at completely aligns to an exonic region of the only FlyBase-annotated transcript isoform of Mad. Gene sequence location is 2L:3146056..3159643.
User Contributed Data
	Beta version, feedback welcome
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	apg: apang Homozygotes when raised at 19 show occasional absence of one or both claws; veins L4 and L5 interrupted; fertile at 19 but become sterile when shifted to 28; produce embryos with range of germ band abnormalities. Homozygous pupal lethal when raised at 28; pharate adults show defective tarsal development of all six legs; condensed, poorly developed and curved metatarsus and tarsi; duplications in tibial and tarsal segments; claws absent. Temperature sensitive period first instar to early pupa.
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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FB2013_03	Controlled Vocabulary Terms A1-7 LL1 motor neuron A1-7 DO2 motor neuron A1-7 VL3/4 motor neuron
FB2013_02
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 23D3-23D3   Limits computationally determined from genome sequence between P{PZ}toc01361 and P{lacW}Madk00237
Experimentally Determined Cytological Location
Cytogenetic map Notes References 23D3-23D4   (determined by in situ hybridisation)   (Spradling et al., 1999) 23D-23D   (determined by in situ hybridisation)   (Roch et al., 1998) 23D4-23D6   (determined by in situ hybridisation)   (Wiersdorff et al., 1996) 23D-23D   (determined by in situ hybridisation)   (Newfeld et al., 1994) 23D3-23D4     (BDGP Project Members, 1994-1999)   The Mad gene is missing in Df(2L)S2590, although the distal break of the cytologically defined deficiency falls in the toc gene.   (Laverty, 1998.12.3)
Experimentally Determined Recombination Data
Location	2-7.7
Left of (cM)	dp (Decotto and Ferguson, 2001)
Right of (cM)	al (Decotto and Ferguson, 2001)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\Mad 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 5.45 Gene model reviewed during 5.46 Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) Mad-RA FBtr0077616  NM_057669   2691   455 Mad-RB FBtr0332372  NM_001273063   3262   525
Additional Transcript Data & Comments
Reported size (kB)	2.6 (northern blot) (Sekelsky et al., 1995)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein Mad-PA   FBpp0077302   50.5   455   7.04     NP_477017   AAF51142 Mad-PB   FBpp0304648   57.9   525   7.20     NP_001259992   AGB92529
Additional Polypeptide Data & Comments
Reported size (kDa)	455 (aa) (Sekelsky et al., 1995)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • SMAD domain, Dwarfin-type (IPR001132) MAD homology 1, Dwarfin-type (IPR003619) SMAD/FHA domain (IPR008984) MAD homology, MH1 (IPR013019) Dwarfin (IPR013790) SMAD domain-like (IPR017855)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA202204   AA202795   AA263717   AA391379   AA392100   AA392101   AA438405   AA441174   AA441225   AA441543   AA539039   AA540100   AA816669   AA816974   AA949669   AA951023   AE014134 AAF51142   AC004422   AE014134 AGB92529   AI454978   AI455013   AI513141   AI513620   AI515225   AI518059   AI546112   AQ034169   BF490999   BH900860   BI163299   BI164256   BI165325   BI169409   BI169926   BI170000   BI171249   BI171948   BI214953   BI216138   BI228682   BI228862   BI229137   BI231585   BI232657   BI237417   BI356488   BI358098   BI363759   BI364700   BI366055   BI367563   BI372600   BI373668   BI482569   BI482847   BI484074   BI484645   BI486251   BI564363   BI573625   BI573626   BI578355   BI578782   BI586072   BI590143   BI593573   BI606000   BI608027   BI609936   BI617114   BI620902   BI626043   BI630583   BI630716   BI631198   BI631410   BP558534   BP558535   BT004845 AAO45201   CK133794   CO177113   CO185834   CO196336   CO261982   CO264134   CO269852   CO270549   CO276466   CO276657   CO277001   CO280776   CO281271   CO285725   CO292357   CO300698   CO307823   CO308745   CO309952   CO311667   CO314365   CO319634   CO324989   CO327030   CO328958   CO332489   CO342069   CO342853   CO342899   CO343001   CO344656   CZ466572   DN154119   EC059808   EC061788   EC066274   EC072968   EC074855   EC074871   EC075069   EC078851   EC079657   EC080398   EC081310   EC083101   EC197522   EC197563   EC199384   EC205280   EC207379   EC212643   EC225047   EC225317   EC229064   EC230084   EC233947   EC236711   EC236734   EC240105   EC246182   EC250343   EC251728   EC252926   EC253661   EC257288   EC257532   EC258588   EC259499   EC259513   EC260174   EV587719   GH739737   GH742764   GH743487   GH767787   GH779755   U10328 AAB60230
UniProtKB/Swiss-Prot	P42003
UniProtKB/TrEMBL
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
in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage 1 -- 3   organism   (Fisher et al., 2012) Comment:maternally deposited embryonic stage 4 -- 6   organism   (Fisher et al., 2012) embryonic stage 11 -- 12   anterior midgut primordium   (Fisher et al., 2012) garland cell primordium   (Fisher et al., 2012) posterior midgut primordium   (Fisher et al., 2012) embryonic stage 13 -- 16   embryonic/larval garland cell   (Fisher et al., 2012) embryonic/larval midgut   (Fisher et al., 2012) northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage -- adult stage     (Sekelsky et al., 1995)
Additional Descriptive Data
	Mad 2.6kb transcripts are detected in all developmental stages but are most abundant in early embryos, pupae, and adult females. A minor slightly smaller transcript is also detected in 0-4hr embryos. (Sekelsky et al., 1995)
Marker for
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization Stage Tissue/Position (including subcellular localization) Reference embryonic stage   organism | ubiquitous   (Newfeld et al., 1996) embryonic stage 5   organism | dorsal   (Srinivasan et al., 2002) Comment:Expressed in a dorsal midline stripe embryonic stage 15   neuron | subset of presumptive embryonic/larval central nervous system   (Rawson et al., 2003) embryonic stage 15 -- 16   embryonic/larval midgut   (Rawson et al., 2003) embryonic stage 16 -- first instar larval stage   RP2 motor neuron   (Rawson et al., 2003) RP3 motor neuron   (Rawson et al., 2003) aCC neuron   (Rawson et al., 2003) U neuron   (Rawson et al., 2003) larval stage   Tv neuron   (Baumgardt et al., 2007) third instar larval stage   germline stem cell   • nucleus (Gilboa and Lehmann, 2004) Comment:in primordial germ cell (PGC), at mid-third instar (ML3, 96hr AEL) embryonic/larval neuromuscular junction | restricted   • type I terminal bouton (Nahm et al., 2010) Comment:phosphorylated Mad (P-Mad) morphogenetic furrow   (Yuva-Aydemir et al., 2011) Comment:antibody specific to phosphorylated Mad eye disc anterior to the morphogenetic furrow | restricted   (Yuva-Aydemir et al., 2011) Comment:antibody specific to phosphorylated Mad embryonic/larval neuromuscular junction   • terminal bouton (Higashi-Kovtun et al., 2010) Comment:antibody specific to phosphorylated Mad (pMad) embryonic/larval neuromuscular junction   • postsynaptic membrane (Higashi-Kovtun et al., 2010) Comment:antibody specific to phosphorylated Mad (pMad) motor neuron of embryonic/larval neuromuscular junction   • nucleus (Higashi-Kovtun et al., 2010) Comment:antibody specific to phosphorylated Mad (pMad) ventral nerve cord   • nucleus (Higashi-Kovtun et al., 2010) Comment:antibody specific to phosphorylated Mad (pMad) wandering third instar larval stage   wing disc | anterior/posterior compartment boundary & gradient   (Bornemann et al., 2004) Comment:antibody specific to phosphorylated Mad adult stage | female   female germline stem cell   (Song et al., 2004) Comment:high expression level cystoblast   (Song et al., 2004) Comment:low expression level adult stage | male   male germline stem cell   (Kawase et al., 2004)
Additional Descriptive Data
	staining was for for pMad (Gilboa and Lehmann, 2004) Phosphorylated Mad protein is distributed in a gradient, with the highest levels near the anterior/posterior compartment boundary. (Bornemann et al., 2004) In female adults, Mad protein is expressed at a high level in germline stem cells, and at a low level in cystoblasts. (Song et al., 2004) Mad protein is ubiquitously distributed in embryos. (Newfeld et al., 1996) Phosphorylated Mad protein is localized to a subset of motor neurons in late stage embryos. (Rawson et al., 2003) Phosphorylated Mad protein is expressed in the presynaptic domain of type I boutons at the neuromuscular junction. (Nahm et al., 2010) In male adults, Mad protein is expressed in germline stem cells. (Kawase et al., 2004) The phosphorylated form of Mad protein is detected in two stripes in the eye disc: one corresponding to the morphogenetic furrow and the other anterior to it. No expression of phosphorylated Mad is detected in glial cells in the eye disc. (Yuva-Aydemir et al., 2011)
Marker for
	male germline stem cell germline stem cell female germline stem cell
Subcellular Localization (GO Cellular Component)
CV term Evidence References
cytoplasm inferred from direct assay (Newfeld et al., 1996, Chen et al., 2006) nucleus inferred from direct assay (Kim et al., 1997, Chen et al., 2006)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools
Reference	See Gelbart and Emmert, 2010.10.13 for analysis details and data files for all genes.
FlyAtlas Anatomy Microarray
FlyAtlas Anatomical Expression Data (FlyAtlas-RNA.adult) (FlyAtlas-RNA.larva)    Styles Linear Logarithmic Heatmap Back-to-back    Scales gene max expression Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Nearly all larval and adult tissues/organs expressed at moderate levels. Expression at high levels in the following post-embryonic organs or tissues: adult ovary. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, larval/adult central nervous system, adult crop, larval/adult midgut, larval/adult hindgut, larval/adult Malpighian tubules, adult heart, larval/adult fat body, larval/adult salivary gland, larval trachea, adult spermathecae, adult male accessory gland, larval/adult carcass. [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 (>999.999) Linear, scaled to maximum expression level Tissue   Expression Level Larval Central Nervous System    231.625 Larval Midgut    336.5 Larval Hindgut    163.7 Larval Malpighian Tubules    203.5 Larval Fat Body    235.5 Larval Salivary Gland    183.4 Larval Trachea    125.625 Larval Carcass    215.9 Adult Head    131.8 Adult Eye    83.7 Adult Brain    153 Adult Thoracic-Abdominal Ganglion    145.5 Adult Crop    146.3 Adult Midgut    279.2 Adult Hindgut    119.9 Adult Malpighian Tubules    127.4 Adult Fat Body    314.5 Adult Salivary Gland    194.6 Adult Heart    170.35 Adult VirginFemale Spermatheca    243.9 Adult InseminatedFemale Spermatheca    248.5 Adult Ovary    524.1 Adult Testis    77.5 Adult Male Accessory Gland    168.2 Adult Carcass    150.5 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    231.625 Larval Midgut    336.5 Larval Hindgut    163.7 Larval Malpighian Tubules    203.5 Larval Fat Body    235.5 Larval Salivary Gland    183.4 Larval Trachea    125.625 Larval Carcass    215.9 Adult Head    131.8 Adult Eye    83.7 Adult Brain    153 Adult Thoracic-Abdominal Ganglion    145.5 Adult Crop    146.3 Adult Midgut    279.2 Adult Hindgut    119.9 Adult Malpighian Tubules    127.4 Adult Fat Body    314.5 Adult Salivary Gland    194.6 Adult Heart    170.35 Adult VirginFemale Spermatheca    243.9 Adult InseminatedFemale Spermatheca    248.5 Adult Ovary  524.1 Adult Testis    77.5 Adult Male Accessory Gland    168.2 Adult Carcass    150.5 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    231.625 Larval Midgut    336.5 Larval Hindgut    163.7 Larval Malpighian Tubules    203.5 Larval Fat Body    235.5 Larval Salivary Gland    183.4 Larval Trachea    125.625 Larval Carcass    215.9 Adult Head    131.8 Adult Eye    83.7 Adult Brain    153 Adult Thoracic-Abdominal Ganglion    145.5 Adult Crop    146.3 Adult Midgut    279.2 Adult Hindgut    119.9 Adult Malpighian Tubules    127.4 Adult Fat Body    314.5 Adult Salivary Gland    194.6 Adult Heart    170.35 Adult VirginFemale Spermatheca    243.9 Adult InseminatedFemale Spermatheca    248.5 Adult Ovary    524.1 Adult Testis    77.5 Adult Male Accessory Gland    168.2 Adult Carcass    150.5 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    231.625 Larval Midgut    336.5 Larval Hindgut    163.7 Larval Malpighian Tubules    203.5 Larval Fat Body    235.5 Larval Salivary Gland    183.4 Larval Trachea    125.625 Larval Carcass    215.9 Adult Head    131.8 Adult Eye    83.7 Adult Brain    153 Adult Thoracic-Abdominal Ganglion    145.5 Adult Crop    146.3 Adult Midgut    279.2 Adult Hindgut    119.9 Adult Malpighian Tubules    127.4 Adult Fat Body    314.5 Adult Salivary Gland    194.6 Adult Heart    170.35 Adult VirginFemale Spermatheca    243.9 Adult InseminatedFemale Spermatheca    248.5 Adult Ovary    524.1 Adult Testis    77.5 Adult Male Accessory Gland    168.2 Adult Carcass    150.5 Expression Level Scale  Very high  log, scaled to maximum expression level Tissue   Expression Level Larval Central Nervous System    231.625 Larval Midgut    336.5 Larval Hindgut    163.7 Larval Malpighian Tubules    203.5 Larval Fat Body    235.5 Larval Salivary Gland    183.4 Larval Trachea    125.625 Larval Carcass    215.9 Adult Head    131.8 Adult Eye    83.7 Adult Brain    153 Adult Thoracic-Abdominal Ganglion    145.5 Adult Crop    146.3 Adult Midgut    279.2 Adult Hindgut    119.9 Adult Malpighian Tubules    127.4 Adult Fat Body    314.5 Adult Salivary Gland    194.6 Adult Heart    170.35 Adult VirginFemale Spermatheca    243.9 Adult InseminatedFemale Spermatheca    248.5 Adult Ovary    524.1 Adult Testis    77.5 Adult Male Accessory Gland    168.2 Adult Carcass    150.5 Expression Level Scale  None   Low   Moderate   High  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    231.625 Larval Midgut    336.5 Larval Hindgut    163.7 Larval Malpighian Tubules    203.5 Larval Fat Body    235.5 Larval Salivary Gland    183.4 Larval Trachea    125.625 Larval Carcass    215.9 Adult Head    131.8 Adult Eye    83.7 Adult Brain    153 Adult Thoracic-Abdominal Ganglion    145.5 Adult Crop    146.3 Adult Midgut    279.2 Adult Hindgut    119.9 Adult Malpighian Tubules    127.4 Adult Fat Body    314.5 Adult Salivary Gland    194.6 Adult Heart    170.35 Adult VirginFemale Spermatheca    243.9 Adult InseminatedFemale Spermatheca    248.5 Adult Ovary  524.1 Adult Testis    77.5 Adult Male Accessory Gland    168.2 Adult Carcass    150.5 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    231.625 Larval Midgut    336.5 Larval Hindgut    163.7 Larval Malpighian Tubules    203.5 Larval Fat Body    235.5 Larval Salivary Gland    183.4 Larval Trachea    125.625 Larval Carcass    215.9 Adult Head    131.8 Adult Eye    83.7 Adult Brain    153 Adult Thoracic-Abdominal Ganglion    145.5 Adult Crop    146.3 Adult Midgut    279.2 Adult Hindgut    119.9 Adult Malpighian Tubules    127.4 Adult Fat Body    314.5 Adult Salivary Gland    194.6 Adult Heart    170.35 Adult VirginFemale Spermatheca    243.9 Adult InseminatedFemale Spermatheca    248.5 Adult Ovary    524.1 Adult Testis    77.5 Adult Male Accessory Gland    168.2 Adult Carcass    150.5 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    231.625 Larval Midgut    336.5 Larval Hindgut    163.7 Larval Malpighian Tubules    203.5 Larval Fat Body    235.5 Larval Salivary Gland    183.4 Larval Trachea    125.625 Larval Carcass    215.9 Adult Head    131.8 Adult Eye    83.7 Adult Brain    153 Adult Thoracic-Abdominal Ganglion    145.5 Adult Crop    146.3 Adult Midgut    279.2 Adult Hindgut    119.9 Adult Malpighian Tubules    127.4 Adult Fat Body    314.5 Adult Salivary Gland    194.6 Adult Heart    170.35 Adult VirginFemale Spermatheca    243.9 Adult InseminatedFemale Spermatheca    248.5 Adult Ovary    524.1 Adult Testis    77.5 Adult Male Accessory Gland    168.2 Adult Carcass    150.5 Expression Level Scale  None   Low   Moderate   High   Very high  Heatmap Tissue   Expression Level Larval Central Nervous System     Larval Midgut     Larval Hindgut     Larval Malpighian Tubules     Larval Fat Body     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     Adult Testis     Adult Male Accessory Gland     Adult Carcass     FlyAtlas Organ/Tissue Expression, larval vs. adult Larval Expression Level Tissue Adult Expression Level   NA  Head    131.8   NA  Eye    83.7   NA  Brain    153   231.625  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    145.5   NA  Crop    146.3   336.5  Midgut    279.2   163.7  Hindgut    119.9   203.5  Malpighian Tubules    127.4   235.5  Fat Body    314.5   183.4  Salivary Gland    194.6   NA  Heart    170.35   125.625  Trachea    NA   NA  VirginFemale Spermatheca    243.9   NA  InseminatedFemale Spermatheca    248.5   NA  Ovary    524.1   NA  Testis    77.5   NA  Male Accessory Gland    168.2   215.9  Carcass    150.5 FlyAtlas Anatomical Expression Data (Chintapalli et al., 2007)
modENCODE Anatomy RNA-Seq
modENCODE Tissue Expression Data (modENCODE_mRNA-Seq_tissues)    Styles Linear Logarithmic Heatmap    Scales gene max expression Moderate expression bin max High expression bin max Extremely high expression bin max [download data (TSV)] Guide to modENCODE expression level colors   No/Extremely low expression (0 - 0)   Very low expression (1 - 3)   Low expression (4 - 10)   Moderate expression (11 - 25)   Moderately high expression (26 - 50)   High expression (51 - 100)   Very high expression (101 - 1000)   Extremely high expression (>1000) Linear, scaled to maximum expression level Tissue   Expression Level imaginal disc, larvae L3 wandering    16 central nervous system, larvae L3    12 central nervous system, pupae P8    11 head, virgin 1-day female    6 head, virgin 4-day female    5 head, virgin 20-day female    8 head, mated 1-day female    6 head, mated 4-day female    6 head, mated 20-day female    6 head, mated 1-day male    8 head, mated 4-day male    10 head, mated 20-day male    11 salivary gland, larvae L3 wandering    3 salivary gland, white prepupae    10 digestive system, larvae L3 wandering    12 digestive system, 1-day adult    9 digestive system, 4-day adult    8 digestive system, 20-day adult    10 fat body, larvae L3 wandering    5 fat body, white prepupae    9 fat body, pupae P8    6 carcass, larvae L3 wandering    11 carcass, 1-day adult    6 carcass, 4-day adult    6 carcass, 20-day adult    6 ovary, virgin 4-day female    35 ovary, mated 4-day female    32 testis, mated 4-day male    9 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to Moderate expression Tissue   Expression Level imaginal disc, larvae L3 wandering    16 central nervous system, larvae L3    12 central nervous system, pupae P8    11 head, virgin 1-day female    6 head, virgin 4-day female    5 head, virgin 20-day female    8 head, mated 1-day female    6 head, mated 4-day female    6 head, mated 20-day female    6 head, mated 1-day male    8 head, mated 4-day male    10 head, mated 20-day male    11 salivary gland, larvae L3 wandering    3 salivary gland, white prepupae    10 digestive system, larvae L3 wandering    12 digestive system, 1-day adult    9 digestive system, 4-day adult    8 digestive system, 20-day adult    10 fat body, larvae L3 wandering    5 fat body, white prepupae    9 fat body, pupae P8    6 carcass, larvae L3 wandering    11 carcass, 1-day adult    6 carcass, 4-day adult    6 carcass, 20-day adult    6 ovary, virgin 4-day female  (35) ovary, mated 4-day female  (32) testis, mated 4-day male    9 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Tissue   Expression Level imaginal disc, larvae L3 wandering    16 central nervous system, larvae L3    12 central nervous system, pupae P8    11 head, virgin 1-day female    6 head, virgin 4-day female    5 head, virgin 20-day female    8 head, mated 1-day female    6 head, mated 4-day female    6 head, mated 20-day female    6 head, mated 1-day male    8 head, mated 4-day male    10 head, mated 20-day male    11 salivary gland, larvae L3 wandering    3 salivary gland, white prepupae    10 digestive system, larvae L3 wandering    12 digestive system, 1-day adult    9 digestive system, 4-day adult    8 digestive system, 20-day adult    10 fat body, larvae L3 wandering    5 fat body, white prepupae    9 fat body, pupae P8    6 carcass, larvae L3 wandering    11 carcass, 1-day adult    6 carcass, 4-day adult    6 carcass, 20-day adult    6 ovary, virgin 4-day female    35 ovary, mated 4-day female    32 testis, mated 4-day male    9 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Tissue   Expression Level imaginal disc, larvae L3 wandering    16 central nervous system, larvae L3    12 central nervous system, pupae P8    11 head, virgin 1-day female    6 head, virgin 4-day female    5 head, virgin 20-day female    8 head, mated 1-day female    6 head, mated 4-day female    6 head, mated 20-day female    6 head, mated 1-day male    8 head, mated 4-day male    10 head, mated 20-day male    11 salivary gland, larvae L3 wandering    3 salivary gland, white prepupae    10 digestive system, larvae L3 wandering    12 digestive system, 1-day adult    9 digestive system, 4-day adult    8 digestive system, 20-day adult    10 fat body, larvae L3 wandering    5 fat body, white prepupae    9 fat body, pupae P8    6 carcass, larvae L3 wandering    11 carcass, 1-day adult    6 carcass, 4-day adult    6 carcass, 20-day adult    6 ovary, virgin 4-day female    35 ovary, mated 4-day female    32 testis, mated 4-day male    9 accessory gland, mated 4-day male    4 Expression Level Scale  Extremely high  log, scaled to maximum expression level Tissue   Expression Level imaginal disc, larvae L3 wandering    16 central nervous system, larvae L3    12 central nervous system, pupae P8    11 head, virgin 1-day female    6 head, virgin 4-day female    5 head, virgin 20-day female    8 head, mated 1-day female    6 head, mated 4-day female    6 head, mated 20-day female    6 head, mated 1-day male    8 head, mated 4-day male    10 head, mated 20-day male    11 salivary gland, larvae L3 wandering    3 salivary gland, white prepupae    10 digestive system, larvae L3 wandering    12 digestive system, 1-day adult    9 digestive system, 4-day adult    8 digestive system, 20-day adult    10 fat body, larvae L3 wandering    5 fat body, white prepupae    9 fat body, pupae P8    6 carcass, larvae L3 wandering    11 carcass, 1-day adult    6 carcass, 4-day adult    6 carcass, 20-day adult    6 ovary, virgin 4-day female    35 ovary, mated 4-day female    32 testis, mated 4-day male    9 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to Moderate expression Tissue   Expression Level imaginal disc, larvae L3 wandering    16 central nervous system, larvae L3    12 central nervous system, pupae P8    11 head, virgin 1-day female    6 head, virgin 4-day female    5 head, virgin 20-day female    8 head, mated 1-day female    6 head, mated 4-day female    6 head, mated 20-day female    6 head, mated 1-day male    8 head, mated 4-day male    10 head, mated 20-day male    11 salivary gland, larvae L3 wandering    3 salivary gland, white prepupae    10 digestive system, larvae L3 wandering    12 digestive system, 1-day adult    9 digestive system, 4-day adult    8 digestive system, 20-day adult    10 fat body, larvae L3 wandering    5 fat body, white prepupae    9 fat body, pupae P8    6 carcass, larvae L3 wandering    11 carcass, 1-day adult    6 carcass, 4-day adult    6 carcass, 20-day adult    6 ovary, virgin 4-day female  (35) ovary, mated 4-day female  32 testis, mated 4-day male    9 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Tissue   Expression Level imaginal disc, larvae L3 wandering    16 central nervous system, larvae L3    12 central nervous system, pupae P8    11 head, virgin 1-day female    6 head, virgin 4-day female    5 head, virgin 20-day female    8 head, mated 1-day female    6 head, mated 4-day female    6 head, mated 20-day female    6 head, mated 1-day male    8 head, mated 4-day male    10 head, mated 20-day male    11 salivary gland, larvae L3 wandering    3 salivary gland, white prepupae    10 digestive system, larvae L3 wandering    12 digestive system, 1-day adult    9 digestive system, 4-day adult    8 digestive system, 20-day adult    10 fat body, larvae L3 wandering    5 fat body, white prepupae    9 fat body, pupae P8    6 carcass, larvae L3 wandering    11 carcass, 1-day adult    6 carcass, 4-day adult    6 carcass, 20-day adult    6 ovary, virgin 4-day female    35 ovary, mated 4-day female    32 testis, mated 4-day male    9 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Tissue   Expression Level imaginal disc, larvae L3 wandering    16 central nervous system, larvae L3    12 central nervous system, pupae P8    11 head, virgin 1-day female    6 head, virgin 4-day female    5 head, virgin 20-day female    8 head, mated 1-day female    6 head, mated 4-day female    6 head, mated 20-day female    6 head, mated 1-day male    8 head, mated 4-day male    10 head, mated 20-day male    11 salivary gland, larvae L3 wandering    3 salivary gland, white prepupae    10 digestive system, larvae L3 wandering    12 digestive system, 1-day adult    9 digestive system, 4-day adult    8 digestive system, 20-day adult    10 fat body, larvae L3 wandering    5 fat body, white prepupae    9 fat body, pupae P8    6 carcass, larvae L3 wandering    11 carcass, 1-day adult    6 carcass, 4-day adult    6 carcass, 20-day adult    6 ovary, virgin 4-day female    35 ovary, mated 4-day female    32 testis, mated 4-day male    9 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Tissue   Expression Level imaginal disc, larvae L3 wandering     central nervous system, larvae L3     central nervous system, pupae P8     head, virgin 1-day female     head, virgin 4-day female     head, virgin 20-day female     head, mated 1-day female     head, mated 4-day female     head, mated 20-day female     head, mated 1-day male     head, mated 4-day male     head, mated 20-day male     salivary gland, larvae L3 wandering     salivary gland, white prepupae     digestive system, larvae L3 wandering     digestive system, 1-day adult     digestive system, 4-day adult     digestive system, 20-day adult     fat body, larvae L3 wandering     fat body, white prepupae     fat body, pupae P8     carcass, larvae L3 wandering     carcass, 1-day adult     carcass, 4-day adult     carcass, 20-day adult     ovary, virgin 4-day female     ovary, mated 4-day female     testis, mated 4-day male     accessory gland, mated 4-day male
modENCODE Development RNA-Seq
modENCODE Temporal Expression Data (modENCODE_mRNA-Seq_U)    Styles Linear Logarithmic Heatmap    Scales gene max expression 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 moderate expression. Peak expression observed at stages throughout embryogenesis. [download data (TSV)] Guide to modENCODE expression level colors   No/Extremely low expression (0 - 0)   Very low expression (1 - 3)   Low expression (4 - 10)   Moderate expression (11 - 25)   Moderately high expression (26 - 50)   High expression (51 - 100)   Very high expression (101 - 1000)   Extremely high expression (>1000) Linear, scaled to maximum expression level Developmental Stage   Expression Level embryo 00-02hr    77 embryo 02-04hr    83 embryo 04-06hr    31 embryo 06-08hr    49 embryo 08-10hr    60 embryo 10-12hr    57 embryo 12-14hr    37 embryo 14-16hr    45 embryo 16-18hr    37 embryo 18-20hr    25 embryo 20-22hr    41 embryo 22-24hr    35 larva L1    9 larva L2    8 larva L3 12hr old    9 larva L3 puffstage 1-2    11 larva L3 puffstage 3-6    21 larva L3 puffstage 7-9    25 white prepupae new    27 white prepupae 12hr    24 white prepupae 24hr    31 pupae 2d postWPP    20 pupae 3d postWPP    15 pupae 4d postWPP    16 adult male 01day    14 adult male 05day    16 adult male 30day    16 adult female 01day    30 adult female 05day    39 adult female 30day    40 Expression Level Scale  Very low   Low   Moderate   Moderately high   High  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (77) embryo 02-04hr  (83) embryo 04-06hr  (31) embryo 06-08hr  (49) embryo 08-10hr  (60) embryo 10-12hr  (57) embryo 12-14hr  (37) embryo 14-16hr  (45) embryo 16-18hr  (37) embryo 18-20hr    25 embryo 20-22hr  (41) embryo 22-24hr  (35) larva L1    9 larva L2    8 larva L3 12hr old    9 larva L3 puffstage 1-2    11 larva L3 puffstage 3-6    21 larva L3 puffstage 7-9    25 white prepupae new  (27) white prepupae 12hr    24 white prepupae 24hr  (31) pupae 2d postWPP    20 pupae 3d postWPP    15 pupae 4d postWPP    16 adult male 01day    14 adult male 05day    16 adult male 30day    16 adult female 01day  (30) adult female 05day  (39) adult female 30day  (40) Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    77 embryo 02-04hr    83 embryo 04-06hr    31 embryo 06-08hr    49 embryo 08-10hr    60 embryo 10-12hr    57 embryo 12-14hr    37 embryo 14-16hr    45 embryo 16-18hr    37 embryo 18-20hr    25 embryo 20-22hr    41 embryo 22-24hr    35 larva L1    9 larva L2    8 larva L3 12hr old    9 larva L3 puffstage 1-2    11 larva L3 puffstage 3-6    21 larva L3 puffstage 7-9    25 white prepupae new    27 white prepupae 12hr    24 white prepupae 24hr    31 pupae 2d postWPP    20 pupae 3d postWPP    15 pupae 4d postWPP    16 adult male 01day    14 adult male 05day    16 adult male 30day    16 adult female 01day    30 adult female 05day    39 adult female 30day    40 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    77 embryo 02-04hr    83 embryo 04-06hr    31 embryo 06-08hr    49 embryo 08-10hr    60 embryo 10-12hr    57 embryo 12-14hr    37 embryo 14-16hr    45 embryo 16-18hr    37 embryo 18-20hr    25 embryo 20-22hr    41 embryo 22-24hr    35 larva L1    9 larva L2    8 larva L3 12hr old    9 larva L3 puffstage 1-2    11 larva L3 puffstage 3-6    21 larva L3 puffstage 7-9    25 white prepupae new    27 white prepupae 12hr    24 white prepupae 24hr    31 pupae 2d postWPP    20 pupae 3d postWPP    15 pupae 4d postWPP    16 adult male 01day    14 adult male 05day    16 adult male 30day    16 adult female 01day    30 adult female 05day    39 adult female 30day    40 Expression Level Scale  Extremely high  log, scaled to maximum expression level Developmental Stage   Expression Level embryo 00-02hr    77 embryo 02-04hr    83 embryo 04-06hr    31 embryo 06-08hr    49 embryo 08-10hr    60 embryo 10-12hr    57 embryo 12-14hr    37 embryo 14-16hr    45 embryo 16-18hr    37 embryo 18-20hr    25 embryo 20-22hr    41 embryo 22-24hr    35 larva L1    9 larva L2    8 larva L3 12hr old    9 larva L3 puffstage 1-2    11 larva L3 puffstage 3-6    21 larva L3 puffstage 7-9    25 white prepupae new    27 white prepupae 12hr    24 white prepupae 24hr    31 pupae 2d postWPP    20 pupae 3d postWPP    15 pupae 4d postWPP    16 adult male 01day    14 adult male 05day    16 adult male 30day    16 adult female 01day    30 adult female 05day    39 adult female 30day    40 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (77) embryo 02-04hr  (83) embryo 04-06hr  31 embryo 06-08hr  (49) embryo 08-10hr  (60) embryo 10-12hr  (57) embryo 12-14hr  (37) embryo 14-16hr  (45) embryo 16-18hr  (37) embryo 18-20hr    25 embryo 20-22hr  (41) embryo 22-24hr  (35) larva L1    9 larva L2    8 larva L3 12hr old    9 larva L3 puffstage 1-2    11 larva L3 puffstage 3-6    21 larva L3 puffstage 7-9    25 white prepupae new  27 white prepupae 12hr    24 white prepupae 24hr  31 pupae 2d postWPP    20 pupae 3d postWPP    15 pupae 4d postWPP    16 adult male 01day    14 adult male 05day    16 adult male 30day    16 adult female 01day  30 adult female 05day  (39) adult female 30day  (40) Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    77 embryo 02-04hr    83 embryo 04-06hr    31 embryo 06-08hr    49 embryo 08-10hr    60 embryo 10-12hr    57 embryo 12-14hr    37 embryo 14-16hr    45 embryo 16-18hr    37 embryo 18-20hr    25 embryo 20-22hr    41 embryo 22-24hr    35 larva L1    9 larva L2    8 larva L3 12hr old    9 larva L3 puffstage 1-2    11 larva L3 puffstage 3-6    21 larva L3 puffstage 7-9    25 white prepupae new    27 white prepupae 12hr    24 white prepupae 24hr    31 pupae 2d postWPP    20 pupae 3d postWPP    15 pupae 4d postWPP    16 adult male 01day    14 adult male 05day    16 adult male 30day    16 adult female 01day    30 adult female 05day    39 adult female 30day    40 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    77 embryo 02-04hr    83 embryo 04-06hr    31 embryo 06-08hr    49 embryo 08-10hr    60 embryo 10-12hr    57 embryo 12-14hr    37 embryo 14-16hr    45 embryo 16-18hr    37 embryo 18-20hr    25 embryo 20-22hr    41 embryo 22-24hr    35 larva L1    9 larva L2    8 larva L3 12hr old    9 larva L3 puffstage 1-2    11 larva L3 puffstage 3-6    21 larva L3 puffstage 7-9    25 white prepupae new    27 white prepupae 12hr    24 white prepupae 24hr    31 pupae 2d postWPP    20 pupae 3d postWPP    15 pupae 4d postWPP    16 adult male 01day    14 adult male 05day    16 adult male 30day    16 adult female 01day    30 adult female 05day    39 adult female 30day    40 Expression Level Scale  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     modENCODE Temporal Expression Data (Graveley et al., 2011)
modENCODE Cell Lines RNA-Seq
modENCODE Cell Line Expression Data (modENCODE_mRNA-Seq_cell.A) (modENCODE_mRNA-Seq_cell.B)    Styles Linear Logarithmic Heatmap    Scales gene max expression Moderate expression bin max High expression bin max Extremely high expression bin max [download data (TSV)] Guide to modENCODE expression level colors   No/Extremely low expression (0 - 0)   Very low expression (1 - 3)   Low expression (4 - 10)   Moderate expression (11 - 25)   Moderately high expression (26 - 50)   High expression (51 - 100)   Very high expression (101 - 1000)   Extremely high expression (>1000) Linear, scaled to maximum expression level Cell Line   Expression Level Schneider line 2 S2R+    11 Schneider line 2 Sg4    12 embryonic 1182-4H    14 embryonic GM2    4 embryonic Kc167    18 embryonic S1    10 embryonic S3    12 leg disc CME L1    8 wing disc CME-W2    7 wing disc ML-DmD8    12 wing disc ML-DmD9    15 wing disc ML-DmD16-c3    18 wing disc ML-DmD21    54 wing disc ML-DmD32    18 haltere disc ML-DmD17-c3    70 eye-antennal disc ML-DmD11    16 antennal disc ML-DmD20-c5    8 mixed discs ML-DmD4-c1    17 CNS ML-DmBG1-c1    13 CNS ML-DmBG2-c2    17 tumorous blood cells mbn2    13 ovary fGS/OSS    25 ovary OSC    19 ovary OSS    14 Expression Level Scale  Very low   Low   Moderate   Moderately high   High  Linear, scaled to Moderate expression Cell Line   Expression Level Schneider line 2 S2R+    11 Schneider line 2 Sg4    12 embryonic 1182-4H    14 embryonic GM2    4 embryonic Kc167    18 embryonic S1    10 embryonic S3    12 leg disc CME L1    8 wing disc CME-W2    7 wing disc ML-DmD8    12 wing disc ML-DmD9    15 wing disc ML-DmD16-c3    18 wing disc ML-DmD21  (54) wing disc ML-DmD32    18 haltere disc ML-DmD17-c3  (70) eye-antennal disc ML-DmD11    16 antennal disc ML-DmD20-c5    8 mixed discs ML-DmD4-c1    17 CNS ML-DmBG1-c1    13 CNS ML-DmBG2-c2    17 tumorous blood cells mbn2    13 ovary fGS/OSS    25 ovary OSC    19 ovary OSS    14 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Cell Line   Expression Level Schneider line 2 S2R+    11 Schneider line 2 Sg4    12 embryonic 1182-4H    14 embryonic GM2    4 embryonic Kc167    18 embryonic S1    10 embryonic S3    12 leg disc CME L1    8 wing disc CME-W2    7 wing disc ML-DmD8    12 wing disc ML-DmD9    15 wing disc ML-DmD16-c3    18 wing disc ML-DmD21    54 wing disc ML-DmD32    18 haltere disc ML-DmD17-c3    70 eye-antennal disc ML-DmD11    16 antennal disc ML-DmD20-c5    8 mixed discs ML-DmD4-c1    17 CNS ML-DmBG1-c1    13 CNS ML-DmBG2-c2    17 tumorous blood cells mbn2    13 ovary fGS/OSS    25 ovary OSC    19 ovary OSS    14 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Cell Line   Expression Level Schneider line 2 S2R+    11 Schneider line 2 Sg4    12 embryonic 1182-4H    14 embryonic GM2    4 embryonic Kc167    18 embryonic S1    10 embryonic S3    12 leg disc CME L1    8 wing disc CME-W2    7 wing disc ML-DmD8    12 wing disc ML-DmD9    15 wing disc ML-DmD16-c3    18 wing disc ML-DmD21    54 wing disc ML-DmD32    18 haltere disc ML-DmD17-c3    70 eye-antennal disc ML-DmD11    16 antennal disc ML-DmD20-c5    8 mixed discs ML-DmD4-c1    17 CNS ML-DmBG1-c1    13 CNS ML-DmBG2-c2    17 tumorous blood cells mbn2    13 ovary fGS/OSS    25 ovary OSC    19 ovary OSS    14 Expression Level Scale  Extremely high  log, scaled to maximum expression level Cell Line   Expression Level Schneider line 2 S2R+    11 Schneider line 2 Sg4    12 embryonic 1182-4H    14 embryonic GM2    4 embryonic Kc167    18 embryonic S1    10 embryonic S3    12 leg disc CME L1    8 wing disc CME-W2    7 wing disc ML-DmD8    12 wing disc ML-DmD9    15 wing disc ML-DmD16-c3    18 wing disc ML-DmD21    54 wing disc ML-DmD32    18 haltere disc ML-DmD17-c3    70 eye-antennal disc ML-DmD11    16 antennal disc ML-DmD20-c5    8 mixed discs ML-DmD4-c1    17 CNS ML-DmBG1-c1    13 CNS ML-DmBG2-c2    17 tumorous blood cells mbn2    13 ovary fGS/OSS    25 ovary OSC    19 ovary OSS    14 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Moderate expression Cell Line   Expression Level Schneider line 2 S2R+    11 Schneider line 2 Sg4    12 embryonic 1182-4H    14 embryonic GM2    4 embryonic Kc167    18 embryonic S1    10 embryonic S3    12 leg disc CME L1    8 wing disc CME-W2    7 wing disc ML-DmD8    12 wing disc ML-DmD9    15 wing disc ML-DmD16-c3    18 wing disc ML-DmD21  (54) wing disc ML-DmD32    18 haltere disc ML-DmD17-c3  (70) eye-antennal disc ML-DmD11    16 antennal disc ML-DmD20-c5    8 mixed discs ML-DmD4-c1    17 CNS ML-DmBG1-c1    13 CNS ML-DmBG2-c2    17 tumorous blood cells mbn2    13 ovary fGS/OSS    25 ovary OSC    19 ovary OSS    14 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Cell Line   Expression Level Schneider line 2 S2R+    11 Schneider line 2 Sg4    12 embryonic 1182-4H    14 embryonic GM2    4 embryonic Kc167    18 embryonic S1    10 embryonic S3    12 leg disc CME L1    8 wing disc CME-W2    7 wing disc ML-DmD8    12 wing disc ML-DmD9    15 wing disc ML-DmD16-c3    18 wing disc ML-DmD21    54 wing disc ML-DmD32    18 haltere disc ML-DmD17-c3    70 eye-antennal disc ML-DmD11    16 antennal disc ML-DmD20-c5    8 mixed discs ML-DmD4-c1    17 CNS ML-DmBG1-c1    13 CNS ML-DmBG2-c2    17 tumorous blood cells mbn2    13 ovary fGS/OSS    25 ovary OSC    19 ovary OSS    14 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Cell Line   Expression Level Schneider line 2 S2R+    11 Schneider line 2 Sg4    12 embryonic 1182-4H    14 embryonic GM2    4 embryonic Kc167    18 embryonic S1    10 embryonic S3    12 leg disc CME L1    8 wing disc CME-W2    7 wing disc ML-DmD8    12 wing disc ML-DmD9    15 wing disc ML-DmD16-c3    18 wing disc ML-DmD21    54 wing disc ML-DmD32    18 haltere disc ML-DmD17-c3    70 eye-antennal disc ML-DmD11    16 antennal disc ML-DmD20-c5    8 mixed discs ML-DmD4-c1    17 CNS ML-DmBG1-c1    13 CNS ML-DmBG2-c2    17 tumorous blood cells mbn2    13 ovary fGS/OSS    25 ovary OSC    19 ovary OSS    14 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Cell Line   Expression Level Schneider line 2 S2R+     Schneider line 2 Sg4     embryonic 1182-4H     embryonic GM2     embryonic Kc167     embryonic S1     embryonic S3     leg disc CME L1     wing disc CME-W2     wing disc ML-DmD8     wing disc ML-DmD9     wing disc ML-DmD16-c3     wing disc ML-DmD21     wing disc ML-DmD32     haltere disc ML-DmD17-c3     eye-antennal disc ML-DmD11     antennal disc ML-DmD20-c5     mixed discs ML-DmD4-c1     CNS ML-DmBG1-c1     CNS ML-DmBG2-c2     tumorous blood cells mbn2     ovary fGS/OSS     ovary OSC     ovary OSS
modENCODE Treatments RNA-Seq
modENCODE Treatment Expression Data (modENCODE_mRNA-Seq_treatments)    Styles Linear Logarithmic Heatmap    Scales gene max expression Moderate expression bin max High expression bin max Extremely high expression bin max [download data (TSV)] Guide to modENCODE expression level colors   No/Extremely low expression (0 - 0)   Very low expression (1 - 3)   Low expression (4 - 10)   Moderate expression (11 - 25)   Moderately high expression (26 - 50)   High expression (51 - 100)   Very high expression (101 - 1000)   Extremely high expression (>1000) Linear, scaled to maximum expression level Treatment   Expression Level extended cold, 4-day adult    14 cold shock, 4-day adult    10 heat shock, 4-day adult    16 Cadmium 50 mM 6 hrs, larvae L3    8 Cadmium 50 mM 12 hrs, larvae L3    8 Cadmium 50 mM 48 hrs, 4-day adult    14 Cadmium 100 mM 48 hrs, 4-day adult    24 Copper 0.5 mM 12 hrs, larvae L3    10 Copper 15 mM 48 hrs, 4-day adult    18 Zinc 5 mM 12 hrs, larvae L3    5 Zinc 4.5 mM 48 hrs, 4-day adult    9 Ethanol 2.5% 3 hrs, larvae L3    30 Ethanol 5% 3 hrs, larvae L3    11 Ethanol 10% 3 hrs, larvae L3    7 Caffeine 1.5 mg/ml 4 hrs, larvae L3    8 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    26 Caffeine 25 mg/ml 48 hrs, 4-day adult    16 Paraquat 5 mM 48 hrs, 4-day adult    24 Paraquat 10 mM 48 hrs, 4-day adult    22 Rotenone 2 μg 12 hrs, larvae L3    7 Rotenone 8 μg 12 hrs, larvae L3    5 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to Moderate expression Treatment   Expression Level extended cold, 4-day adult    14 cold shock, 4-day adult    10 heat shock, 4-day adult    16 Cadmium 50 mM 6 hrs, larvae L3    8 Cadmium 50 mM 12 hrs, larvae L3    8 Cadmium 50 mM 48 hrs, 4-day adult    14 Cadmium 100 mM 48 hrs, 4-day adult    24 Copper 0.5 mM 12 hrs, larvae L3    10 Copper 15 mM 48 hrs, 4-day adult    18 Zinc 5 mM 12 hrs, larvae L3    5 Zinc 4.5 mM 48 hrs, 4-day adult    9 Ethanol 2.5% 3 hrs, larvae L3  (30) Ethanol 5% 3 hrs, larvae L3    11 Ethanol 10% 3 hrs, larvae L3    7 Caffeine 1.5 mg/ml 4 hrs, larvae L3    8 Caffeine 2.5 mg/ml 48 hrs, 4-day adult  26 Caffeine 25 mg/ml 48 hrs, 4-day adult    16 Paraquat 5 mM 48 hrs, 4-day adult    24 Paraquat 10 mM 48 hrs, 4-day adult    22 Rotenone 2 μg 12 hrs, larvae L3    7 Rotenone 8 μg 12 hrs, larvae L3    5 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Treatment   Expression Level extended cold, 4-day adult    14 cold shock, 4-day adult    10 heat shock, 4-day adult    16 Cadmium 50 mM 6 hrs, larvae L3    8 Cadmium 50 mM 12 hrs, larvae L3    8 Cadmium 50 mM 48 hrs, 4-day adult    14 Cadmium 100 mM 48 hrs, 4-day adult    24 Copper 0.5 mM 12 hrs, larvae L3    10 Copper 15 mM 48 hrs, 4-day adult    18 Zinc 5 mM 12 hrs, larvae L3    5 Zinc 4.5 mM 48 hrs, 4-day adult    9 Ethanol 2.5% 3 hrs, larvae L3    30 Ethanol 5% 3 hrs, larvae L3    11 Ethanol 10% 3 hrs, larvae L3    7 Caffeine 1.5 mg/ml 4 hrs, larvae L3    8 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    26 Caffeine 25 mg/ml 48 hrs, 4-day adult    16 Paraquat 5 mM 48 hrs, 4-day adult    24 Paraquat 10 mM 48 hrs, 4-day adult    22 Rotenone 2 μg 12 hrs, larvae L3    7 Rotenone 8 μg 12 hrs, larvae L3    5 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Treatment   Expression Level extended cold, 4-day adult    14 cold shock, 4-day adult    10 heat shock, 4-day adult    16 Cadmium 50 mM 6 hrs, larvae L3    8 Cadmium 50 mM 12 hrs, larvae L3    8 Cadmium 50 mM 48 hrs, 4-day adult    14 Cadmium 100 mM 48 hrs, 4-day adult    24 Copper 0.5 mM 12 hrs, larvae L3    10 Copper 15 mM 48 hrs, 4-day adult    18 Zinc 5 mM 12 hrs, larvae L3    5 Zinc 4.5 mM 48 hrs, 4-day adult    9 Ethanol 2.5% 3 hrs, larvae L3    30 Ethanol 5% 3 hrs, larvae L3    11 Ethanol 10% 3 hrs, larvae L3    7 Caffeine 1.5 mg/ml 4 hrs, larvae L3    8 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    26 Caffeine 25 mg/ml 48 hrs, 4-day adult    16 Paraquat 5 mM 48 hrs, 4-day adult    24 Paraquat 10 mM 48 hrs, 4-day adult    22 Rotenone 2 μg 12 hrs, larvae L3    7 Rotenone 8 μg 12 hrs, larvae L3    5 Expression Level Scale  Extremely high  log, scaled to maximum expression level Treatment   Expression Level extended cold, 4-day adult    14 cold shock, 4-day adult    10 heat shock, 4-day adult    16 Cadmium 50 mM 6 hrs, larvae L3    8 Cadmium 50 mM 12 hrs, larvae L3    8 Cadmium 50 mM 48 hrs, 4-day adult    14 Cadmium 100 mM 48 hrs, 4-day adult    24 Copper 0.5 mM 12 hrs, larvae L3    10 Copper 15 mM 48 hrs, 4-day adult    18 Zinc 5 mM 12 hrs, larvae L3    5 Zinc 4.5 mM 48 hrs, 4-day adult    9 Ethanol 2.5% 3 hrs, larvae L3    30 Ethanol 5% 3 hrs, larvae L3    11 Ethanol 10% 3 hrs, larvae L3    7 Caffeine 1.5 mg/ml 4 hrs, larvae L3    8 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    26 Caffeine 25 mg/ml 48 hrs, 4-day adult    16 Paraquat 5 mM 48 hrs, 4-day adult    24 Paraquat 10 mM 48 hrs, 4-day adult    22 Rotenone 2 μg 12 hrs, larvae L3    7 Rotenone 8 μg 12 hrs, larvae L3    5 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to Moderate expression Treatment   Expression Level extended cold, 4-day adult    14 cold shock, 4-day adult    10 heat shock, 4-day adult    16 Cadmium 50 mM 6 hrs, larvae L3    8 Cadmium 50 mM 12 hrs, larvae L3    8 Cadmium 50 mM 48 hrs, 4-day adult    14 Cadmium 100 mM 48 hrs, 4-day adult    24 Copper 0.5 mM 12 hrs, larvae L3    10 Copper 15 mM 48 hrs, 4-day adult    18 Zinc 5 mM 12 hrs, larvae L3    5 Zinc 4.5 mM 48 hrs, 4-day adult    9 Ethanol 2.5% 3 hrs, larvae L3  30 Ethanol 5% 3 hrs, larvae L3    11 Ethanol 10% 3 hrs, larvae L3    7 Caffeine 1.5 mg/ml 4 hrs, larvae L3    8 Caffeine 2.5 mg/ml 48 hrs, 4-day adult  26 Caffeine 25 mg/ml 48 hrs, 4-day adult    16 Paraquat 5 mM 48 hrs, 4-day adult    24 Paraquat 10 mM 48 hrs, 4-day adult    22 Rotenone 2 μg 12 hrs, larvae L3    7 Rotenone 8 μg 12 hrs, larvae L3    5 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Treatment   Expression Level extended cold, 4-day adult    14 cold shock, 4-day adult    10 heat shock, 4-day adult    16 Cadmium 50 mM 6 hrs, larvae L3    8 Cadmium 50 mM 12 hrs, larvae L3    8 Cadmium 50 mM 48 hrs, 4-day adult    14 Cadmium 100 mM 48 hrs, 4-day adult    24 Copper 0.5 mM 12 hrs, larvae L3    10 Copper 15 mM 48 hrs, 4-day adult    18 Zinc 5 mM 12 hrs, larvae L3    5 Zinc 4.5 mM 48 hrs, 4-day adult    9 Ethanol 2.5% 3 hrs, larvae L3    30 Ethanol 5% 3 hrs, larvae L3    11 Ethanol 10% 3 hrs, larvae L3    7 Caffeine 1.5 mg/ml 4 hrs, larvae L3    8 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    26 Caffeine 25 mg/ml 48 hrs, 4-day adult    16 Paraquat 5 mM 48 hrs, 4-day adult    24 Paraquat 10 mM 48 hrs, 4-day adult    22 Rotenone 2 μg 12 hrs, larvae L3    7 Rotenone 8 μg 12 hrs, larvae L3    5 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Treatment   Expression Level extended cold, 4-day adult    14 cold shock, 4-day adult    10 heat shock, 4-day adult    16 Cadmium 50 mM 6 hrs, larvae L3    8 Cadmium 50 mM 12 hrs, larvae L3    8 Cadmium 50 mM 48 hrs, 4-day adult    14 Cadmium 100 mM 48 hrs, 4-day adult    24 Copper 0.5 mM 12 hrs, larvae L3    10 Copper 15 mM 48 hrs, 4-day adult    18 Zinc 5 mM 12 hrs, larvae L3    5 Zinc 4.5 mM 48 hrs, 4-day adult    9 Ethanol 2.5% 3 hrs, larvae L3    30 Ethanol 5% 3 hrs, larvae L3    11 Ethanol 10% 3 hrs, larvae L3    7 Caffeine 1.5 mg/ml 4 hrs, larvae L3    8 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    26 Caffeine 25 mg/ml 48 hrs, 4-day adult    16 Paraquat 5 mM 48 hrs, 4-day adult    24 Paraquat 10 mM 48 hrs, 4-day adult    22 Rotenone 2 μg 12 hrs, larvae L3    7 Rotenone 8 μg 12 hrs, larvae L3    5 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Treatment   Expression Level extended cold, 4-day adult     cold shock, 4-day adult     heat shock, 4-day adult     Cadmium 50 mM 6 hrs, larvae L3     Cadmium 50 mM 12 hrs, larvae L3     Cadmium 50 mM 48 hrs, 4-day adult     Cadmium 100 mM 48 hrs, 4-day adult     Copper 0.5 mM 12 hrs, larvae L3     Copper 15 mM 48 hrs, 4-day adult     Zinc 5 mM 12 hrs, larvae L3     Zinc 4.5 mM 48 hrs, 4-day adult     Ethanol 2.5% 3 hrs, larvae L3     Ethanol 5% 3 hrs, larvae L3     Ethanol 10% 3 hrs, larvae L3     Caffeine 1.5 mg/ml 4 hrs, larvae L3     Caffeine 2.5 mg/ml 48 hrs, 4-day adult     Caffeine 25 mg/ml 48 hrs, 4-day adult     Paraquat 5 mM 48 hrs, 4-day adult     Paraquat 10 mM 48 hrs, 4-day adult     Rotenone 2 μg 12 hrs, larvae L3     Rotenone 8 μg 12 hrs, larvae L3
Expression Clusters
	A cluster of genes with similar mRNA expression dynamics across development. (The modENCODE Consortium, 2010, Graveley et al., 2011) mE2_34_mRNA_expression_cluster_17 mE1_20_mRNA_expression_cluster_03
External Data & Images
Linkouts	BDGP expression data • RE72705 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0011648 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG12399-RA FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for Mad Stages(s) 1-3 Stages(s) 4-6 Stages(s) 9-10 Stages(s) 11-12
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele cell lethal | recessive | somatic clone MadB1 lethal (with Mad2) Madk00223 lethal (with MadB1) Madk00223 lethal (with Madk00223) Mad2 MadB1 lethal | embryonic stage | germline clone | recessive | rescuable maternal effect Mad2L-23-1 Mad2L-112-1 Mad2L-136-1 Mad2L-163-23 lethal | embryonic stage | maternal effect | recessive Mad12 MadEz lethal | larval stage | recessive MadB1 lethal | pupal stage, with Scer\GAL4da.G32 MaddsRNA.Scer\UAS.cEa lethal | pupal stage, with Scer\GAL4pnr-MD237 MadGD4121 lethal | pupal stage | recessive Mad1 Mad7 Mad10 Mad12 lethal | pupal stage | recessive | heat sensitive Madapg lethal | recessive Mad1-1 Mad1-2 Mad4-1 Mad5 Mad6-2 Mad6 Mad7-1 Mad8-2 Mad8 Mad9-2 Mad9 Mad11 Mad17-3 Mad18-3 Mad20-2 Mad22-3 MadD3 MadD14 MadD15 MadD24 Madk00223 Madk00237 Madk05807a MadKG00581 MadP1 partially lethal - majority live, with Scer\GAL4Dll-md23 Mad1.Scer\UAS partially lethal - majority live, with Scer\GAL4how-24B Mad1.Scer\UAS partially lethal - majority live, with Scer\GAL4T80 Mad1.Scer\UAS viable MadEz viable (with Mad2L-112-1) Mad6 viable (with Mad2L-136-1) Mad6 viable (with Mad2L-163-23) Mad6 viable (with Mad6) Mad2L-112-1 Mad2L-136-1 Mad2L-163-23 Sterility Allele female sterile (with Mad2L-112-1) Mad6 female sterile (with Mad2L-136-1) Mad6 female sterile (with Mad2L-163-23) Mad6 female sterile (with Mad6) Mad2L-112-1 Mad2L-136-1 Mad2L-163-23 fertile MadEz sterile | recessive | heat sensitive Madapg Other Phenotypes Allele decreased cell number | adult stage | conditional, with Scer\GAL4nos.UTR.T:Hsim\VP16 MadmiRNA.2XartmiR.Scer\UAS.P\T decreased cell number | adult stage | somatic clone Mad12 decreased cell number | somatic clone Mad12 decreased cell number | third instar larval stage, with Scer\GAL4repo.PU MadKK109050 increased cell death | third instar larval stage, with Scer\GAL4ap-md544 MadNIG.12399R mitotic cell cycle defective | cell autonomous | somatic clone Mad12 neuroanatomy defective Madunspecified neuroanatomy defective (with Df(2L)C28) Mad1 neuroanatomy defective (with Mad1) MadB65 neuroanatomy defective (with MadB65) Mad1 neuroanatomy defective | third instar larval stage Madk00237 neuroanatomy defective | third instar larval stage (with Mad1) Madk00237 neuroanatomy defective | third instar larval stage (with Madk00237) Mad1 neurophysiology defective (with Mad1) Mad12 neurophysiology defective (with Mad12) Mad1 neurophysiology defective | third instar larval stage Madk00237 size defective, with Scer\GAL4A9 MadScer\UAS.cNa size defective, with Scer\GAL4vg.MQ.Exel MaddsRNA.Scer\UAS.cPa small body | larval stage (with Mad1) MadB65 small body | larval stage (with MadB65) Mad1 visible, with Scer\GAL4A9 MaddsRNA.Scer\UAS.cEa MadMGM.Scer\UAS MadMMM.Scer\UAS MadSDVD.Scer\UAS visible, with Scer\GAL4ap-md544 MadMGM.Scer\UAS MadMMM.Scer\UAS visible, with Scer\GAL4Bx-MS1096-KE MaddsRNA.Scer\UAS.cEa visible, with Scer\GAL4ptc-559.1 MadMGM.Scer\UAS MadMMM.Scer\UAS MadScer\UAS.cNa visible, with Scer\GAL4sca-537.4 MaddsRNA.Scer\UAS.cUa MadMGM.Scer\UAS MadScer\UAS.cNa visible, with Scer\GAL4sd-SG29.1 MadGM8.Scer\UAS MadMGM.Scer\UAS MadMMM.Scer\UAS visible, with Scer\GAL4vg.int2.1 MadScer\UAS.cNa visible, with Scer\GAL4vg.PM MadScer\UAS.cNa visible | dominant Madk00237 MadKG00581 visible | embryonic stage, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 MaddsRNA.Scer\UAS.P\T MadMGM.Scer\UAS visible | heat sensitive (with Madapg) Madk00237 visible | heat sensitive (with Madk00237) Madapg visible | recessive Madapg MadD16 MadP1 visible | recessive | somatic clone Mad1-2 visible | somatic clone Mad12 visible | somatic clone, with Scer\GAL4Act5C.PI MaddsRNA.Scer\UAS.cEa MadMGM.Scer\UAS wild-type Mad+t11.5 MadUbi-p63E.T:Hsap\MYC Phenotype manifest in Allele abdominal segment Madapg adult midgut precursor cell | somatic clone Mad12 anterior wing margin | somatic clone, with Scer\GAL4Act5C.PI MaddsRNA.Scer\UAS.cEa bouton (with Df(2L)C28) Mad1 crossvein, with Scer\GAL469B Mad1.Scer\UAS crossvein | ectopic, with Scer\GAL4ptc-559.1 MadMMM.Scer\UAS MadMGM.Scer\UAS crystal cell Mad12 denticle belt Mad10 Mad12 MadcHa denticle belt, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 MadMGM.Scer\UAS denticle belt | ectopic, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 MaddsRNA.Scer\UAS.P\T dorsal triple row | ectopic, with Scer\GAL4sd-SG29.1 MadMMM.Scer\UAS MadMGM.Scer\UAS dorsal triple row | supernumerary, with Scer\GAL4A9 MadMGM.Scer\UAS dorsal triple row | supernumerary, with Scer\GAL4sd-SG29.1 MadMGM.Scer\UAS egg Mad12 egg chamber Mad12 embryonic/first instar larval cuticle Mad10 Mad12 embryonic/first instar larval cuticle | maternal effect Mad12 MadEz embryonic/first instar larval cuticle | ventral MadcHa embryonic/larval dorsal branch Mad12 MadB1 embryonic/larval fat body MadB1 embryonic/larval midgut | anterior MadB1 embryonic/larval neuromuscular junction | third instar larval stage Madk00237 embryonic/larval neuromuscular junction | third instar larval stage (with Mad1) Madk00237 embryonic/larval neuromuscular junction | third instar larval stage (with Madk00237) Mad1 embryonic/larval optic lobe MadB1 embryonic/larval somatic muscle (with Mad1) MadB65 embryonic/larval somatic muscle (with MadB65) Mad1 embryonic/larval tracheal system Mad12 embryonic/larval ventral anastomosis Mad12 eye Mad1-2 Mad9-2 eye | somatic clone MadB1 eye-antennal disc Mad1-1 Mad1-2 Mad12 Mad17-3 Mad18-3 Mad20-2 Mad22-3 Mad4-1 Mad6-2 Mad7-1 Mad8-2 Mad9-2 eye disc | cell autonomous | somatic clone Mad12 eye disc | posterior | somatic clone Mad1-2 MadB1 eye disc | third instar larval stage, with Scer\GAL4repo.PU MadKK109050 fat body (with Mad7) Mad12 fat body (with Mad12) Mad7 female germline stem cell Mad12 Mad9 female germline stem cell, with Scer\GAL4nos.UTR.T:Hsim\VP16 MadmiRNA.2XartmiR.Scer\UAS.P\T female germline stem cell | adult stage | somatic clone Mad12 female germline stem cell | somatic clone Mad12 frons | ectopic | somatic clone Mad1-2 gastric caecum Mad12 germarium Mad12 germarium, with Scer\GAL4nos.UTR.T:Hsim\VP16 MadmiRNA.2XartmiR.Scer\UAS.P\T histoblast | somatic clone Mad1 histoblast nest | somatic clone Mad1 imaginal disc MadB1 Madk00223 imaginal disc | maternal effect Mad12 MadEz interommatidial bristle | pupal stage P7 | somatic clone Mad12 leg Mad1-1 Mad9-2 leg | distal | heat sensitive (with Madapg) Madk00237 leg | distal | heat sensitive (with Madk00237) Madapg leg | ectopic, with Scer\GAL4ptc-559.1 MadScer\UAS.cNa leg | heat sensitive (with Madapg) Madk00237 leg | heat sensitive (with Madk00237) Madapg leg | somatic clone Mad12 male germline stem cell | germline clone Mad12 medial triple row | supernumerary, with Scer\GAL4A9 MadMGM.Scer\UAS medial triple row | supernumerary, with Scer\GAL4sd-SG29.1 MadMGM.Scer\UAS morphogenetic furrow Mad1-2 Mad9-2 neuromuscular junction, with Scer\GAL4Mhc.PW MadScer\UAS.T:Avic\GFP-EGFP neuromuscular junction (with Df(2L)C28) Mad1 neuromuscular junction (with Mad1) MadB65 neuromuscular junction (with MadB65) Mad1 NMJ bouton | third instar larval stage Madk00237 NMJ bouton | third instar larval stage (with Mad1) Madk00237 NMJ bouton | third instar larval stage (with Madk00237) Mad1 ommatidium | somatic clone Mad1-2 ovariole Mad12 ovary, with Scer\GAL4nos.UTR.T:Hsim\VP16 MadmiRNA.2XartmiR.Scer\UAS.P\T photoreceptor cell Mad1-2 Mad9-2 photoreceptor cell | cell non-autonomous | somatic clone MadB1 photoreceptor cell | somatic clone Mad1-2 MadB1 pleurum | somatic clone Mad12 posterior crossvein Mad6 secondary pigment cell | pupal stage P7 | somatic clone Mad12 synapse Madunspecified tarsal segment Madapg tarsal segment | heat sensitive Madapg tarsal segment | somatic clone Mad12 t-bar Madunspecified tergite | somatic clone Mad12 tertiary pigment cell | pupal stage P7 | somatic clone Mad12 testis | germline clone MadB1 tibia | heat sensitive Madapg wing Mad1-1 Mad9-2 MadKG00581 MadScer\UAS.cNa Madk00237 MadSDVD.Scer\UAS wing, with Scer\GAL469B Mad1.Scer\UAS wing, with Scer\GAL4nub-AC-62 MadScer\UAS.cNa wing, with Scer\GAL4ptc-559.1 MadScer\UAS.cNa wing, with Scer\GAL4sca-537.4 MaddsRNA.Scer\UAS.cUa wing, with Scer\GAL4T80 MadScer\UAS.cNa wing, with Scer\GAL4vg.int2.1 MadScer\UAS.cNa wing, with Scer\GAL4vg.PM MadScer\UAS.cNa wing | temperature conditional, with Scer\GAL4Bx-MS1096-KE MaddsRNA.Scer\UAS.cEa wing blade, with Scer\GAL4vg.MQ.Exel MaddsRNA.Scer\UAS.cPa wing cell MadSDVD.Scer\UAS wing disc MadScer\UAS.cNa wing disc | somatic clone Mad12 MadB1 wing disc | third instar larval stage, with Scer\GAL4ap-md544 MadNIG.12399R wing margin, with Scer\GAL469B Mad1.Scer\UAS wing margin (with Mad11) Mad12 wing margin (with Mad12) Mad11 wing margin | somatic clone, with Scer\GAL4Act5C.PI MadMGM.Scer\UAS wing margin bristle, with Scer\GAL4A9 MaddsRNA.Scer\UAS.cEa wing margin bristle | ectopic, with Scer\GAL4sca-537.4 MadMGM.Scer\UAS MaddsRNA.Scer\UAS.cUa wing margin bristle | ectopic, with Scer\GAL4sd-SG29.1 MadGM8.Scer\UAS wing pouch, with Scer\GAL4sd-SG29.1 MadMGM.Scer\UAS wing sensillum | ectopic, with Scer\GAL4A9 MadMMM.Scer\UAS wing sensillum | ectopic, with Scer\GAL4sca-537.4 MadMGM.Scer\UAS MaddsRNA.Scer\UAS.cUa wing vein Mad6 MadScer\UAS.cNa MadSDVD.Scer\UAS wing vein, with Scer\GAL469B Mad1.Scer\UAS wing vein, with Scer\GAL471B MadScer\UAS.cUa wing vein, with Scer\GAL4A9 MadSDVD.Scer\UAS wing vein, with Scer\GAL4Bx-MS1096-KE MaddsRNA.Scer\UAS.cEa wing vein, with Scer\GAL4nub-AC-62 MadScer\UAS.cNa wing vein, with Scer\GAL4sca-537.4 MaddsRNA.Scer\UAS.cUa wing vein, with Scer\GAL4vg.MQ.Exel MaddsRNA.Scer\UAS.cPa wing vein | ectopic, with Scer\GAL471B MadScer\UAS.cUa wing vein | ectopic, with Scer\GAL4A9 MadScer\UAS.cNa wing vein | ectopic, with Scer\GAL4ptc-559.1 MadScer\UAS.cNa MadMMM.Scer\UAS MadMGM.Scer\UAS wing vein | ectopic, with Scer\GAL4sca-537.4 MadScer\UAS.cNa MadMGM.Scer\UAS wing vein | ectopic, with Scer\GAL4vg.int2.1 MadScer\UAS.cNa wing vein L2 (with Mad11) Mad12 wing vein L2 (with Mad12) Mad11 wing vein L2 | cell autonomous | somatic clone Mad12 wing vein L3 | cell autonomous | somatic clone Mad12 wing vein L4 Madapg wing vein L4 (with Mad11) Mad12 wing vein L4 (with Mad12) Mad11 wing vein L5 Mad6 Madapg wing vein L5 (with Mad11) Mad12 wing vein L5 (with Mad12) Mad11
Classical Alleles ( 49 )
For All Classical Alleles Show
Allele of Mad Class Mutagen Stocks Known lesion Madk00237 P-element activity 2 -- Mad1-2 hypomorphic allele - genetic evidence P-element activity 1 Yes Mad8-2 hypomorphic allele - genetic evidence P-element activity 1 Yes Madc00574 piggyBac transposase 1 -- Madk05807a P-element activity 1 -- MadKG00581 P-element activity 1 -- MadMI02462 1 -- Mad10 amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes Mad12 amorphic allele - genetic evidence, loss of function allele ethyl methanesulfonate 0 Yes Mad1 amorphic allele - genetic evidence, gain of function allele ethyl methanesulfonate 0 Yes Mad7 amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes Mad1-1 hypomorphic allele - genetic evidence P-element activity 0 Yes Mad11 ethyl methanesulfonate 0 Yes Mad17-3 hypomorphic allele - genetic evidence P-element activity 0 Yes Mad18-3 hypomorphic allele - genetic evidence P-element activity 0 Yes Mad20-2 hypomorphic allele - genetic evidence P-element activity 0 Yes Mad22-3 hypomorphic allele - genetic evidence P-element activity 0 Yes Mad2 loss of function allele ethyl methanesulfonate 0 Yes Mad2L-112-1 ethyl methanesulfonate 0 -- Mad2L-136-1 ethyl methanesulfonate 0 -- Mad2L-163-23 ethyl methanesulfonate 0 -- Mad2L-23-1 ethyl methanesulfonate 0 -- Mad3 loss of function allele ethyl methanesulfonate 0 Yes Mad4-1 hypomorphic allele - genetic evidence P-element activity 0 Yes Mad4 loss of function allele ethyl methanesulfonate 0 Yes Mad5 ethyl methanesulfonate 0 Yes Mad6-2 hypomorphic allele - genetic evidence P-element activity 0 Yes Mad6 ethyl methanesulfonate 0 Yes Mad7-1 hypomorphic allele - genetic evidence P-element activity 0 Yes Mad8 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes Mad9-2 hypomorphic allele - genetic evidence P-element activity 0 Yes Mad9 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes Madapg ethyl methanesulfonate 0 -- MadB1 P-element activity 0 Yes MadB65 0 -- MadD14 ethyl nitrosourea 0 Yes MadD15 ethyl nitrosourea 0 -- MadD16 ethyl nitrosourea 0 Yes MadD24 ethyl nitrosourea 0 Yes MadD3 ethyl nitrosourea 0 -- MadE21 0 -- MadE5 0 -- MadES1 ethyl methanesulfonate 0 -- MadEz hypomorphic allele - genetic evidence ethyl methanesulfonate 0 -- Madk00223 P-element activity 0 Yes MadP-23D P-element activity 0 -- MadP1 P-element activity 0 -- MadSH1449 0 -- Madunspecified 0 --
Alleles Carried on Transgenic Constructs ( 42 )
For All Alleles Carried on Transgenic Constructs Show
Allele of Mad Class Mutagen Stocks Known lesion MadGD4121 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes MadGL01527 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes MadGLV21013 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes MadJF01263 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes MadJF01264 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes MadT:Avic\GFP-SF,T:Zzzz\FLAG in vitro construct - coding region fusion 1 Yes Mad+t11.5 in vitro construct - genomic fragment 0 Yes Mad1.Scer\UAS in vitro construct - regulatory fusion 0 Yes Mad12.Scer\UAS in vitro construct - regulatory fusion 0 Yes Mad4-16 in vitro construct - genomic fragment 0 Yes MadAct5C.T:SV5\V5,T:Zzzz\His6,T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes MadAct5C.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes MadcHa in vitro construct 0 Yes MaddsRNA.cMa in vitro construct - RNAi 0 Yes MaddsRNA.Scer\UAS.cEa in vitro construct - regulatory fusion in vitro construct - RNAi 0 Yes MaddsRNA.Scer\UAS.cEb in vitro construct - regulatory fusion in vitro construct - RNAi 0 Yes MaddsRNA.Scer\UAS.cPa in vitro construct - regulatory fusion in vitro construct - RNAi 0 Yes MaddsRNA.Scer\UAS.cUa in vitro construct - regulatory fusion in vitro construct - RNAi 0 Yes MaddsRNA.Scer\UAS.P\T in vitro construct - regulatory fusion in vitro construct - RNAi 0 Yes MadGM8.Scer\UAS in vitro construct - regulatory fusion in vitro construct - amino acid replacement 0 Yes Madhs.PK in vitro construct - regulatory fusion 0 Yes MadKK109050 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes Madmex1.PN in vitro construct - regulatory fusion 0 Yes MadMGM.Scer\UAS in vitro construct - regulatory fusion in vitro construct - site directed mutagenesis 0 Yes MadmiRNA.2XartmiR.Scer\UAS.P\T in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes MadMMM.Scer\UAS in vitro construct - regulatory fusion in vitro construct - site directed mutagenesis 0 Yes MadMtnA.PP in vitro construct - regulatory fusion 0 Yes MadMtnA.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes MadN.T:Zzzz\FLAG in vitro construct - coding region fusion 0 Yes MadNIG.12399R in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes MadS25A.Scer\UAS.T:Hsap\MYC in vitro construct - amino acid replacement 0 Yes MadS25A.Scer\UAS in vitro construct - regulatory fusion in vitro construct - amino acid replacement 0 Yes MadScer\UAS.cEa in vitro construct - regulatory fusion 0 Yes MadScer\UAS.cNa in vitro construct - regulatory fusion 0 Yes MadScer\UAS.cUa in vitro construct - regulatory fusion 0 Yes MadScer\UAS.T:Avic\GFP-EGFP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes MadScer\UAS.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes MadScer\UAS.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes MadSDVD.Scer\UAS in vitro construct - regulatory fusion in vitro construct - site directed mutagenesis 0 Yes MadT312V.hs in vitro construct - regulatory fusion 0 Yes MadUbi-p63E.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes MadαTub84B.PM in vitro construct - regulatory fusion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2L)BSC28 (Deal and Cook, 2003.1.30) Df(2L)BSC162 (Christensen and Cook, 2006.8.29) Df(2L)BSC163 (Christensen and Cook, 2006.8.29) Df(2L)BSC164 (Christensen and Cook, 2006.8.29) Df(2L)BSC232 (Christensen and Cook, 2007.3.22) Df(2L)BSC238 (Christensen and Cook, 2006.12.5) Df(2L)C28 (Hudson et al., 1998, Raftery et al., 1995, Sekelsky et al., 1995, Newfeld et al., 1996, Hoodless et al., 1996, Wisotzkey et al., 2003) Df(2L)Exel7015 (Ryder, 2004.4.26) Df(2L)JS17 (Yu et al., 1998, Spradling et al., 1999, Su et al., 2001, BDGP Project Members, 1994-1999, Raftery et al., 1995, Sekelsky et al., 1995, Newfeld et al., 1996, Newfeld et al., 1997) Df(2L)JS31   Df(2L)S2590 (BDGP Project Members, 1994-1999) In(2LR)DTD16LDTD42R (Spradling et al., 1999, BDGP Project Members, 1994-1999) Ts(2Rt;3Lt)DTD62+Ts(2Lt;3Rt)HVD7 (Su et al., 2001, Sekelsky et al., 1995, Gelbart, 1996.3.6)
Not disrupted in	Df(2L)C144 (Spradling et al., 1999, Su et al., 2001, BDGP Project Members, 1994-1999, Sekelsky et al., 1995) Df(2L)DTD52-D51 (Su et al., 2001) Df(2L)JS7 (Su et al., 2001, Sekelsky et al., 1995) Df(2L)JS32   Df(2R)E3363 (BDGP Project Members, 1994-1999) T(2;3)DTD46.4 (Su et al., 2001)
Duplicated in	Dp(2;3)JS20 (Sekelsky et al., 1995)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD4121} NA P{KK109050} NA P{TRiP.GL01527} NA P{TRiP.GLV21013} NA P{TRiP.JF01263} NA P{TRiP.JF01264} NA P{UAS-Mad.1} NA P{UAS-Mad.12} NA P{UAS-Mad.FLAG} NA P{UAS-Mad.GM8} NA P{UAS-Mad.MGM} NA P{UAS-Mad.MMM} NA P{UAS-Mad.MWT} NA P{UAS-Mad.N} NA P{UAS-Mad.RNAi.226-807} NA P{UAS-Mad.RNAi.657-1230} NA P{UAS-Mad.RNAi.U} NA P{UAS-Mad.RNAi} NA P{UAS-Mad.S25A} NA P{UAS-Mad.SDVD} NA P{UAS-Mad.U} NA P{UAS-Mad-EGFP} NA P{UAS-Myc-Mad.S25A} NA P{UAS-Myc-Mad} NA P{UASp-Mad.miRNA.2XartmiR} NA P{UASp-Mad.RNAi} NA reporter construct P{Mad.Med.2xsd-lacZ} No P{Mad.Med.2xSu(H)-lacZ} No heat-shock construct P{hs-Mad.K} NA P{hs-Mad.T312V} NA characterization construct P{Mad0-11} NA P{Mad4-16} NA P{mex1Mad} NA P{NIG.12399R} NA P{UmMad} NA P{αTub84B-Mad.M} NA PBac{Mad-GFP.FLAG} NA
Insertions
	Type of insertions Name Expression data insertion of enhancer trap P{lacW}l(2)SH0431SH0431 No P{lacW}MadB1 No P{lacW}Madk00223 No P{lacW}Madk00237 No P{lacW}Madk05807a No P{lacW}MadP-23D No P{lacW}MadSH1449 No miscellaneous insertions Mi{MIC}MadMI02462 NA P{?lacW}Mad1-1 NA P{?lacW}Mad1-2 NA P{?lacW}Mad4-1 NA P{?lacW}Mad6-2 NA P{?lacW}Mad7-1 NA P{?lacW}Mad8-2 NA P{?lacW}Mad9-2 NA P{?lacW}Mad17-3 NA P{?lacW}Mad18-3 NA P{?lacW}Mad20-2 NA P{?lacW}Mad22-3 NA P{SUPor-P}MadKG00581 NA P{walter}MadP1 NA PBac{PB}Madc00574 NA
Gene Ontology: Function, Process & Cellular Component ( 34 unique terms )
Terms Based on Experimental Evidence ( 26 terms )
Molecular Function
CV term Evidence References
protein binding inferred from physical interaction with Med (Wisotzkey et al., 1998) inferred from physical interaction with vis (Hyman et al., 2003) inferred from physical interaction with UniProtKB:Q9V853 (assigned by UniProtKB) (Podos et al., 2001, Liang et al., 2003) RNA polymerase II activating transcription factor binding inferred from physical interaction with shn (Dai et al., 2000) RNA polymerase II distal enhancer sequence-specific DNA binding inferred from direct assay (Xu et al., 1998) RNA polymerase II distal enhancer sequence-specific DNA binding transcription factor activity inferred from direct assay (Kim et al., 1997) RNA polymerase II transcription coactivator activity inferred from direct assay (Dai et al., 2000) sequence-specific DNA binding inferred from direct assay (Lin et al., 2006, Grienenberger et al., 2003, Weiss et al., 2010)
Biological Process
CV term Evidence References
BMP signaling pathway involved in Malpighian tubule cell chemotaxis inferred from direct assay (Bunt et al., 2010) compound eye morphogenesis inferred from mutant phenotype (Cordero et al., 2007) germ-line stem cell division inferred from mutant phenotype (Xie and Spradling, 1998) germ-line stem cell maintenance inferred from mutant phenotype (Xie and Spradling, 1998) histoblast morphogenesis inferred from mutant phenotype (Ninov et al., 2010) imaginal disc-derived leg morphogenesis inferred from mutant phenotype (Marquez et al., 2001) imaginal disc-derived wing morphogenesis inferred from mutant phenotype (Dworkin and Gibson, 2006) imaginal disc-derived wing vein morphogenesis inferred from mutant phenotype (Marquez et al., 2001) negative regulation of gene expression inferred from direct assay (Anderson et al., 2006) negative regulation of S phase of mitotic cell cycle inferred from mutant phenotype (Firth and Baker, 2005) ovarian follicle cell development inferred from mutant phenotype (Kirilly et al., 2005) positive regulation of synaptic growth at neuromuscular junction inferred from mutant phenotype (McCabe et al., 2004, Merino et al., 2009) positive regulation of transcription from RNA polymerase II promoter inferred from expression pattern (Kim et al., 1997) inferred from mutant phenotype (Grienenberger et al., 2003, Xu et al., 1998) regulation of cell differentiation inferred from mutant phenotype (Mathur et al., 2010) somatic stem cell maintenance inferred from mutant phenotype (Kirilly et al., 2005) transforming growth factor beta receptor signaling pathway inferred from genetic interaction with dpp AND inferred from genetic interaction with tkv AND inferred from genetic interaction with wg (Wiersdorff et al., 1996) trunk segmentation inferred from mutant phenotype (Eivers et al., 2009) wing disc anterior/posterior pattern formation inferred from mutant phenotype (Shen and Dahmann, 2005)
Cellular Component
CV term Evidence References
cytoplasm inferred from direct assay (Newfeld et al., 1996, Chen et al., 2006) nucleus inferred from direct assay (Kim et al., 1997, Chen et al., 2006)
Terms Based on Predictions or Assertions ( 13 terms )
Molecular Function
CV term Evidence References
DNA binding traceable author statement (Kawabata and Miyazono, 1999) transforming growth factor beta receptor, pathway-specific cytoplasmic mediator activity non-traceable author statement (Kawabata and Miyazono, 1999)
Biological Process
CV term Evidence References
dorsal closure non-traceable author statement (Harden, 2002) germ-line stem cell division traceable author statement (Lin, 2002) heart development traceable author statement (Cripps and Olson, 2002) negative regulation of salivary gland boundary specification traceable author statement (Bradley et al., 2001) open tracheal system development traceable author statement (Ghabrial et al., 2003) ovarian follicle cell development traceable author statement (Dobens and Raftery, 2000) regulation of transcription from RNA polymerase II promoter non-traceable author statement (Affolter et al., 2001) transforming growth factor beta receptor signaling pathway non-traceable author statement (Swiss-Prot Project Members, 1995.6.1) traceable author statement (Lall and Patel, 2001)
Cellular Component
CV term Evidence References
cytoplasm traceable author statement (Kawabata and Miyazono, 1999) nucleus traceable author statement (Kawabata and Miyazono, 1999, Affolter et al., 2001) transcription factor complex inferred from electronic annotation with InterPro:IPR013019 (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
Mad - Arpc2 anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) Mad - Mtr3 anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) Mad - Tm1 anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) Mad - arm anti tag coimmunoprecipitation, anti tag western blot (Eivers et al., 2011) Mad - mRpL44 anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) Mad - pan anti tag coimmunoprecipitation, anti tag western blot (Eivers et al., 2011)
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions Mad allele Gene References Mad1-2 eya (Curtiss and Mlodzik, 2000) smo (Curtiss and Mlodzik, 2000) Mad1 dpp (Sekelsky et al., 1995) tkv (Hoodless et al., 1996) wit (McCabe et al., 2004) Mad2 dpp (Raftery et al., 1995) Mad3 dpp (Raftery et al., 1995) Mad4 dpp (Raftery et al., 1995) Mad6 tkv (Hoodless et al., 1996) vri (George and Terracol, 1997) Mad10 dpp (Sekelsky et al., 1995) sax (Das et al., 1998) tkv (Das et al., 1998) Mad11 dpp (Sekelsky et al., 1995) Mad12 brk (Shen and Dahmann, 2005) dpp (Sekelsky et al., 1995, Das et al., 1998) sax (Das et al., 1998, Waltzer and Bienz, 1999) sog (Decotto and Ferguson, 2001) tkv (Hoodless et al., 1996, Das et al., 1998) MadB1 brk (Shen and Dahmann, 2005) bsk (Shen and Dahmann, 2005) sca (Wiersdorff et al., 1996) tkv (Wiersdorff et al., 1996) MadD3 dpp (Chen et al., 1998) Med (Chen et al., 1998) put (Chen et al., 1998) shn (Chen et al., 1998) tkv (Chen et al., 1998) MadD14 dpp (Chen et al., 1998) put (Chen et al., 1998) sax (Chen et al., 1998) tkv (Chen et al., 1998) MadD15 dpp (Chen et al., 1998) Med (Chen et al., 1998) put (Chen et al., 1998) shn (Chen et al., 1998) tkv (Chen et al., 1998) MadD16 sax (Chen et al., 1998) tkv (Chen et al., 1998) MadD24 dpp (Chen et al., 1998) Med (Chen et al., 1998) put (Chen et al., 1998) shn (Chen et al., 1998) tkv (Chen et al., 1998) MadE5 vvl (Certel and Johnson, 1998) MadES1 sog (Decotto and Ferguson, 2001) MadEz zen (Hudson et al., 1998) Madk00237 os (Bach et al., 2003) tor (Li and Li, 2003) MadP-23D tkv (Hoodless et al., 1996) MadScer\UAS.cNa Dad (Tsuneizumi et al., 1997) MadUbi-p63E.T:Hsap\MYC dpp (Wiersdorff et al., 1996) unspecified CycE (Horsfield et al., 1998) dally (Kirkpatrick et al., 2006) dpp (Sekelsky et al., 1995) Med (Hudson et al., 1998) nej (Kumar et al., 2004) Pka-C1 (Wiersdorff et al., 1996) Snoo (Takaesu et al., 2006) tkv (Bach et al., 2003)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 59745 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0011648 DroID - A comprehensive database of gene and protein interactions. • FBgn0011648 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 33529 SignaLink - A protein-protein and protein-miRNA interaction database with multi-layered structure and multiple confidence scores. • P42003
Orthologs
OrthoDB Orthologs (66) - based on analysis using Dmel annotation version 5.41
OrthoDB Ortholog Groups
	OrthoDB orthology group searches (Waterhouse et al., 2011, Nucleic Acids Res. 39(Database issue): D283--D288) Drosophila inclusive ortholog search EOG6RBR4C Dipteran inclusive ortholog search EOG6W3SWG Insect inclusive ortholog search EOG6N8Q14 Arthropod inclusive ortholog search EOG6GB5NH Metazoa inclusive ortholog search EOG6X3VV6
Orthologs in Drosophila Species (EOG6RBR4C)
Organism Common Name Gene AAA Syntenic Ortholog Multiple Dmel Genes in this Orthologous Group Drosophila melanogaster fruit fly  Dmel\Mad     Drosophila simulans   Dsim\GD22774 Y   Drosophila sechellia   Dsec\GM18164 Y   Drosophila erecta   Dere\GG24457 Y   Drosophila yakuba   Dyak\GE14935 Y   Drosophila ananassae   Dana\GF20577 Y   Drosophila pseudoobscura pseudoobscura   Dpse\GA28973 Y   Drosophila persimilis   Dper\GL26531 Y   Drosophila willistoni   Dwil\GK18268 Y   Drosophila virilis   Dvir\GJ19553 Y   Drosophila mojavensis   Dmoj\GI17981 Y   Drosophila grimshawi   Dgri\GH10284 Y
Orthologs in non-Drosophila Dipterans (EOG6W3SWG)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Aedes aegypti Yellow fever mosquito  Aaeg\AAEL005513   Anopheles gambiae Malaria mosquito  Agam\Mad   Culex quinquefasciatus Southern house mosquito  Cqui\CPIJ000420
Orthologs in non-Dipteran Insects (EOG6N8Q14)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Apis mellifera Western honey bee  Amel\Ame.16140   Nasonia vitripennis Parasitic wasp  Nvit\Nasvi2EG007038   Nasonia vitripennis Parasitic wasp  Nvit\Nasvi2EG012380   Acromyrmex echinatior Panamanian leafcutter ant  Aech\AECH11334   Atta cephalotes Leafcutter ant  Acep\ACEP16085   Camponotus floridanus Florida carpenter ant  Cflo\CFLO20740   Harpegnathos saltator Jerdons jumping ant  Hsal\HSAL16176   Linepithema humile Argentine ant  Lhum\LH12024   Pogonomyrmex barbatus Red harvester ant  Pbar\PB27356   Pogonomyrmex barbatus Red harvester ant  Pbar\PB27358   Solenopsis invicta Red fire ant  Sinv\SINV11024   Acyrthosiphon pisum Pea aphid  Apim\ACYPI007984   Acyrthosiphon pisum Pea aphid  Apim\ACYPI010124   Bombyx mori Silkmoth  Bmor\BGIBMGA010481   Pediculus humanus Human body louse  Phum\PHUM612490   Pediculus humanus Human body louse  Phum\PHUM612590   Tribolium castaneum Red flour beetle  Tcas\Mad   Tribolium castaneum Red flour beetle  Tcas\GLEAN_14921
Orthologs in non-Insect Arthropods (EOG6GB5NH)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Daphnia pulex Water flea  Dpux\ARHGEF9   Ixodes scapularis Deer tick  Isca\ISCW010869
Orthologs in non-Arthropod Metazoa (EOG6X3VV6)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Caenorhabditis elegans Nematode  Cele\sma-2 Y Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_023107gn Y Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_020722gn Y Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_017642gn Y Danio rerio Zebrafish  Drer\smad5 Y Danio rerio Zebrafish  Drer\smad1 Y Danio rerio Zebrafish  Drer\smad3b Y Danio rerio Zebrafish  Drer\smad9 Y Danio rerio Zebrafish  Drer\OTTDARG00000027522 Y Danio rerio Zebrafish  Drer\smad3a Y Xenopus tropicalis Western clawed frog  Xtro\smad2 Y Xenopus tropicalis Western clawed frog  Xtro\smad9 Y Xenopus tropicalis Western clawed frog  Xtro\smad3 Y Xenopus tropicalis Western clawed frog  Xtro\ENSXETG00000023689 Y Xenopus tropicalis Western clawed frog  Xtro\smad1 Y Gallus gallus Domestic chicken  Ggal\SMAD1 Y Gallus gallus Domestic chicken  Ggal\ENSGALG00000014184 Y Gallus gallus Domestic chicken  Ggal\SMAD5 Y Gallus gallus Domestic chicken  Ggal\SMAD9 Y Gallus gallus Domestic chicken  Ggal\SMAD3 Y Gallus gallus Domestic chicken  Ggal\SMAD2 Y Mus musculus House mouse  Mmus\Smad3 Y Mus musculus House mouse  Mmus\Smad2 Y Mus musculus House mouse  Mmus\Smad9 Y Rattus norvegicus Norway rat  Rnor\Smad3 Y Rattus norvegicus Norway rat  Rnor\Smad9 Y Rattus norvegicus Norway rat  Rnor\Smad5 Y Rattus norvegicus Norway rat  Rnor\ENSRNOG00000018483 Y Rattus norvegicus Norway rat  Rnor\Smad2 Y Homo sapiens Human  Hsap\SMAD2 Y Homo sapiens Human  Hsap\SMAD9 Y
Human Orthologs (2)
Gene OMIM HGNC Hsap\SMAD2     Hsap\SMAD9
AAA Orthologs (11) based on analysis using Dmel annotation version 4.3
Organism Gene Drosophila simulans Dsim\GD22774 Drosophila sechellia Dsec\GM18164 Drosophila erecta Dere\GG24457 Drosophila yakuba Dyak\GE14935 Drosophila ananassae Dana\GF20577 Drosophila pseudoobscura pseudoobscura Dpse\GA28973 Drosophila persimilis Dper\GL26531 Drosophila willistoni Dwil\GK18268 Drosophila virilis Dvir\GJ19553 Drosophila mojavensis Dmoj\GI17981 Drosophila grimshawi Dgri\GH10284
Stocks & Reagents
Stocks Listed in FlyBase ( 14 )
Bloomington	38663 w1118; PBac{Mad-GFP.FLAG}VK00033 7323 w*; Mad1-2 P{neoFRT}40A/CyO 7324 w*; Mad8-2 P{neoFRT}40A/CyO 31315 y1 v1; P{TRiP.JF01263}attP2 31316 y1 v1; P{TRiP.JF01264}attP2 35987 y1 w*; Mi{MIC}MadMI02462 10474 y1 w67c23; P{lacW}Madk00237/CyO 43183 y1 sc* v1; P{TRiP.GL01527}attP40 14578 y1; P{SUPor-P}MadKG00581/CyO; ry506
Harvard	- PBac{PB}Madc00574
Kyoto	102416 y1 w67c23 ; P{lacW}Madk05807a / CyO
VDRC	v12635 w1118; P{GD4121}v12635
	More stocks available...
Genomic Clones ( 2 )
	BACR11E09 BACR48B06 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 132 )
	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	RE72705
Other clones	DPiM_RE72705
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	AT27647 LD01806 LD02325 LD03112 LD03233 LD04033 LD06880 LD10262 LD12458 LD12459 LD12679 LD15979 LD16075 LD16519 LD18741 LD19461 LD21919 LD29643 LD31273 LD37469 LD41967 LD45328 LD45821 LD46958 RE02660 RE03903 RE05188 RE10576 RE11242 RE11334 RE12974 RE13850 RE21119 RE22465 RE23064 RE26306 RE26516 RE26855 RE28758 RE33643 RE38484 RE42655 RE44769 RE48694 RE49867 RE51494 RE53356 RE59603 RE61027 RE65275 RE65622 RE67168 RE67780 RE69655 RE72221 RH03724 RH11683 RH12155 RH14787 RH25810 RH37256 RH47347 RH52159 RH59855 RH60004 RH60727 RH60976 RH66569 RH72061
Other clones	1049 41623 42815 59031 74549 97648 101239 102658 175164 214219.64 214309.83 214311.71 214539.76 276611 CK01.10D.F10 CK01.66C.G7 CK01.77C.G1 CK01.78A.F3 CK01.111D.F5 CK01.129D.F1 CK01.145B.F1 CK01.147C.A10 Dmel_neb_010853_0147_1491 EK003126 EK012853 EK052528 EK060525 EK083308 EK096121 EK099308 EK120933 EK121756 EK122607 EK137602 EK154832 EK156333 EK179205 EK182932 EK195330 EK199921 EK220022 EK225741 EK235939 EK255803 EK259237 EK279650 EK291738 EK303630 EP05527 EP13745 EP15625 EP15713 EP15912 EP19660 FGM315E09 RH09191 RP001019307 RP001039818 RP001046174 RP001054275 RP001055167
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0011648 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 33529
Antibody Information
	polyclonal (Newfeld et al., 1996, Eivers et al., 2009, Nahm et al., 2010, Fuentealba et al., 2008, Chen and Xu, 2010, Colozza et al., 2011, Kaneko et al., 2011, Peterson et al., 2012) polyclonal antibody (Song et al., 2004, Kawase et al., 2004, Bornemann et al., 2004)
Other Information
Discoverer
Etymology
	The gene is named "apang" after the mutant phenotype ("apang" stands for an individual with mutilated and underdeveloped legs in Hindi). (Shekaran and Sharma, 1983)
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of	Source for merge of: E(zen)2 Mad (Ferguson, 1998.8.7) Source for merge of: Mad apg (Galindo et al., 2002) Source for merge of: Mad l(2)k00237 (Laverty, 1998.12.3)
Additional comments
Other Comments
	DNA-protein interactions: genome-wide binding profile assayed for Mad protein in stage 9-11 embryos; ArrayExpress accession number E-MTAB-1184. (Junion et al., 2012) Phosphorylation of Mad protein in the linker region by sgg regulates the development of sensory organs in the anterior-dorsal quadrant of the wing. (Quijano et al., 2011) DNA-protein interactions: genome-wide binding profile assayed for Mad protein in 2-3 hr embryos; see BDTNP1_TFBS_Mad collection report. (MacArthur et al., 2009) The Mad product negatively regulates dac expression in the embryonic head. (Anderson et al., 2006) dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology. (Kiger et al., 2003) The brk silencer serves as a direct target for a protein complex consisting of Mad/Med and shn. (Muller et al., 2003) Hsap\MADH1 and Hsap\MADH4, as well as Mad and Med, can stimulate dpp signalling in limb development. (Marquez et al., 2001) Overexpression of Mad or Hsap\MADH1 in the wing and leg causes a similar phenotype. (Marquez et al., 2001) One of five genes identified as encoding downstream components of the dpp signalling cascade which is necessary for blocking salivary gland gene activation by Scr in the dorsal region of parasegment 2. Mad function is required to block salivary gland formation in dorsal regions of PS2. (Henderson et al., 1999) Mad may recruit nej to effect the transcriptional activation of dpp responsive genes during development. (Waltzer and Bienz, 1999) Loss of function alleles of tkv, put, Mad, Med and shn suppress the CycEJP mutant eye phenotype in combination with dppd-ho. (Horsfield et al., 1998) Mutants show no interaction with Df(2R)Pcl11B or Df(3L)66C-G28. (Nicholls and Gelbart, 1998) Mutants isolated in a screen of the second chromosome identifying genes affecting disc morphology. (Roch et al., 1998) Mad mediated nuclear translocation is essential for Med function. (Wisotzkey et al., 1998) Transcriptional activation of Ubx is subject to competition between dpp-activated Mad and another Smad whose function as a transcriptional repressor depends on high wg signalling. (Yu et al., 1998) The amino-terminal domain of the Mad protein contains a sequence-specific DNA-binding activity that becomes apparent when carboxy-terminal residues are removed. Mad protein binds to and is required for the activation of an enhancer within the vg gene in cells across the entire developing wing blade. (Kim et al., 1997) Mad protein functions downstream of the serine-threonine kinase activity of the tkv receptor. (Newfeld et al., 1997) Signalling by constitutively active tkv mutation is suppressed by heterozygosity for Mad mutations. These results indicate that Mad functions downstream of the tkv receptor. (Hoodless et al., 1996) Mad is required for any response of the visceral mesoderm or endoderm to dpp signals from the visceral mesoderm and is required specifically in cells responding to dpp. Mad can function in the signalling pathway of BMP-4 in Xenopus embryos, and is thus a highly conserved and essential element of the dpp signalling pathway. (Newfeld et al., 1996) Mad is required for dpp signalling during eye development. Clonal analysis demonstrated that this requirement is cell autonomous. Mad is an essential component of the signal transduction pathway downstream of the dpp receptors in responding cells. Mad-mediated dpp signaling is absolutely required for the initiation of the morphogenetic furrow in the eye, but has only a minor role in its subsequent propagation across the eye disc. (Wiersdorff et al., 1996) Dominant enhancer of zen. (Hudson et al., 1995) Mad has been identified independently in two screens, one for dominant enhancers of the dpp mutant phenotype and dosage sensitive interactions with dpp. Mad mutant phenotypes show patterning defects that resemble many dpp mutant phenotypes. Studies of ectopic expression, tissue-specific expression and in situ hybridizations are consistent with a role for Mad downstream of dpp in the signalling pathway. (Newfeld et al., 1995) Genetic characterisation indicates Mad encodes a product essential for dpp function. Molecular analysis demonstrates the Mad protein is a member of a novel protein family that is highly conserved throughout metazoans. (Sekelsky et al., 1995) A genetic enhancer of dpp. (Jackson and Hoffmann, 1994) Loss of function mutations of Mad are dominant maternal effect enhancers of dpp during early embryogenesis and dominant zygotic enhancers of dpp in imaginal discs. (Newfeld et al., 1994)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_057669 NM_001273063 RefSeq (Proteins) • NP_477017 NP_001259992 DDBJ / EMBL / GenBank • AA202795 AAB60230 AAF51142 AAO45201 AC004422 AGB92529 AQ034169 BH900860 BI578782 BT004845 CZ466572 U10328 Entrez Gene - A searchable database of RefSeq genes. • 33529 UniProtKB/Swiss-Prot • P42003
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • SMAD domain, Dwarfin-type (IPR001132) MAD homology 1, Dwarfin-type (IPR003619) SMAD/FHA domain (IPR008984) MAD homology, MH1 (IPR013019) Dwarfin (IPR013790) SMAD domain-like (IPR017855) BDGP expression data - Patterns of gene expression in Drosophila embryogenesis • RE72705
Linkouts
	BioGRID - A database of protein and genetic interactions • 59745 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0011648 DroID - A comprehensive database of gene and protein interactions. • FBgn0011648 DRSC - Results from RNAi screens. • FBgn0011648 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0011648 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG12399-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0011648 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 33529 Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /newgene/mothadpp.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 33529 modMine - Data generated by the modENCODE project. • FBgn0011648 REDfly - A database of transcriptional regulatory elements. • FBgn0011648 SignaLink - A protein-protein and protein-miRNA interaction database with multi-layered structure and multiple confidence scores. • P42003
Synonyms & Secondary IDs ( 38 )
Reported As
Symbol Synonym	2/23 (Roch et al., 1998) apg   c28 (Wisotzkey et al., 2003) CG12399 (Brandt, 2006, Ni et al., 2008, Keleman et al., 2009.8.5, Ni et al., 2011.7.19, Ni et al., 2010.12.1) dMAD (Kang and Bier, 2010) dMad (Hyman et al., 2003) E(zen)2   En(vvl)   l(2)K00237 (Grammont et al., 1997) l(2)k00237   Mad (Tien et al., 2008, Baumgardt et al., 2007, Lin et al., 2006, Pyrowolakis et al., 2004, Gao et al., 2005, Xi and Xie, 2005, Jones et al., 2006, Tyler et al., 2007, Pistillo and Desplan, 2007, Ballard and Wharton, 2007, Yakoby et al., 2007, Liang et al., 2007, Gerlitz et al., 2007, Dudu, 2006, Umulis et al., 2006, Horabin, 2005, Bangi and Wharton, 2006, Layden et al., 2006, Joshi et al., 2006, Bangi and Wharton, 2006, Akasaka et al., 2006, Christensen and Cook, 2006.8.29, Newfeld et al., 2007, Dudu et al., 2006, Crickmore and Mann, 2006, Crickmore and Mann, 2006, Dworkin and Gibson, 2006, Takaesu et al., 2006, Ramel et al., 2007, Christensen and Cook, 2006.8.29, Christensen and Cook, 2006.12.5, Christensen and Cook, 2007.3.22, Christensen and Cook, 2006.8.29, Sun et al., 2012, Ueyama et al., 2008, de Velasco et al., 2006, Shcherbata et al., 2007, Shirinian et al., 2007, Gonzales-Gaitan, 2007, Knoblich, 2006, Bokel et al., 2006, Bokel et al., 2006, de Navas et al., 2006, Gao and Laughon, 2006, Takaesu et al., 2005, Niki et al., 2006, Niki, 2006, Anderson et al., 2006, Kirkpatrick et al., 2006, Sotillos and de Celis, 2006, Mizutani et al., 2005, Stathopoulos and Levine, 2005, Yao et al., 2006, Chen and Schupbach, 2006, Aerts et al., 2007, Ball et al., 2010, Zhu et al., 2012, Kirilly et al., 2005, Fromental-Ramain et al., 2008, Rawson et al., 2003, Mizutani et al., 2006, Herrero et al., 2007, O'Keefe et al., 2011, Yuva-Aydemir et al., 2011, Crickmore and Mann, 2007, Letizia et al., 2007, Newfeld et al., 2008, Campbell and Moser, 2008, Warrior et al., 2008, Shcherbata et al., 2008, Casas-Tinto et al., 2008, Erives et al., 2008, van der Plas et al., 2006, Yamashita et al., 2005, Marques, 2005, Shen et al., 2008, Fernández et al., 2007, Bernardi et al., 2006, Davidson and Erwin, 2006, Wang and Ward, 2010, Lecuit and Le Goff, 2007, Christoforou et al., 2008, Kyoda et al., 2000, Sage et al., 2005, Philippakis et al., 2006, Mikhaylova and Nurminsky, 2011, Firth and Baker, 2005, Hufnagel et al., 2007, Bolivar et al., 2006, Boylan et al., 2008, Jekely and Rorth, 2003, Ables and Drummond-Barbosa, 2010, Ogiso et al., 2011, Sato et al., 2010, Miller et al., 2011, Ben-Zvi et al., 2011, Li et al., 2011, Yao et al., 2008, Nahm et al., 2010, Giorgianni and Mann, 2011, Layalle et al., 2011, Umemori et al., 2009, Oh and Irvine, 2011, Shcherbata et al., 2007, Johnson et al., 2007, Estella et al., 2008, Serpe et al., 2008, Walsh and Carroll, 2007, Xu et al., 2007, Wang et al., 2008, Tyler and Baker, 2007, Kim et al., 2007, Cash and Andrews, 2012, Chang et al., 2008, Yakoby et al., 2008, López-Onieva et al., 2008, Estella and Mann, 2008, Bollenbach et al., 2008, Yao et al., 2008, Tran and Doe, 2008, Ng, 2008, Schwank et al., 2008, Barrio et al., 2007, Li et al., 2007, Umemori et al., 2007, Tanaka-Matakatsu and Du, 2008, Takaesu et al., 2008, Casas-Tinto et al., 2008, Vogler and Urban, 2008, Miguel-Aliaga et al., 2008, Fuentealba et al., 2008, Zeng et al., 2008, Manjon et al., 2007, Li and Li, 2006, Qian et al., 2011, Miles et al., 2008, Sellin et al., 2009, Firth and Baker, 2009, Lilja et al., 2007, Wisotzkey et al., 2003, Pinto et al., 2008, Liu et al., 2009, Korolchuk et al., 2007, Wang et al., 2008, Evans et al., 2009, McKay et al., 2009, Rhiner et al., 2009, Lince-Faria et al., 2009, Lembong et al., 2009, Escudero and Freeman, 2007, Zahedi et al., 2008, Eade and Allan, 2009, Pentek et al., 2009, Kim and Marqués, 2012, Künnapuu et al., 2009, Kamiya et al., 2008, Saller and Bienz, 2001, Zeng et al., 2007, Vrailas and Moses, 2006, Smith-Bolton et al., 2009, Makhijani et al., 2007, Lim et al., 2008, Baena-Lopez et al., 2008, Karim et al., 2012, MacArthur et al., 2009, Brandt, 2006, Grienenberger et al., 2003, Cordero et al., 2007, Garces and Thor, 2006, Guo and Wang, 2009, Oishi et al., 2006, Jensen et al., 2009, Brás-Pereira and Casares, 2008, Glavic et al., 2009, Yu et al., 2009, Eivers et al., 2009, Merino et al., 2009, Mathur et al., 2010, Firth et al., 2010, Neely et al., 2010, Ballard et al., 2010, Wagner et al., 2010, Stefancsik and Sarkar, 2003, Ellis et al., 2010, Zhao et al., 2008, Salzer et al., 2010, Li et al., 2010, Sander et al., 2010, Spokony and White, 2012.5.22, Ishikawa et al., 2010, Twombly et al., 2009, Quijano et al., 2010, Kagey et al., 2012, Koh et al., 2004, Sun et al., 2010, Akiyama et al., 2012, Yagi et al., 2010, Takaesu et al., 2012, Clark et al., 2011, Losada-Pérez et al., 2010, Liu et al., 2010, Padash-Barmchi et al., 2010, Su et al., 2010, Beltran et al., 2007, Ambrus et al., 2007, Weiss et al., 2010, Chen et al., 2010, Eivers et al., 2011, Rodal et al., 2011, König et al., 2011, Liu et al., 2011, Wang et al., 2009, Peluso et al., 2011, Hao et al., 2008, Gao and Laughon, 2007, Rodriguez, 2011, Fischer et al., 2012, Abruzzi et al., 2011, Gomez et al., 2012, Bhattacharya and Baker, 2011, Peterson et al., 2012, Le and Wharton, 2012, Quijano et al., 2011, Dejima et al., 2011, Beck et al., 2012, Gancz et al., 2011, Lembong et al., 2008, Pennetier et al., 2012, Xia et al., 2012, Agelopoulos et al., 2012, Aboukhalil and Bulyk, 2012, Chen et al., 2012, Dahal et al., 2012, Stinchfield et al., 2012) mad (Dworkin and Gibson., 2006, Pappu et al., 2005, Gibson and Perrimon, 2005, Dunlop et al., 2004, Kirilly et al., 2005, Kalinovsky and Scheiffele, 2004, Li et al., 2005, Luschnig et al., 2004, Luschnig et al., 2004, Shivdasani and Ingham, 2003, Zhu and Xie, 2003, Deng and Lin, 2001, Marty et al., 2000, Kawase and Xie, 2003, Kirilly et al., 2003, Bessa et al., 2002, McCabe et al., 2002, Dorfman and Shilo, 2001, Affolter, 2000, Jordan et al., 2000, Zhang et al., 1999, Lin, 1998, Yu et al., 1998, Horsfield et al., 1998, Cho and Blitz, 1998, Xie and Spradling, 1998, Certel and Johnson, 1998, Henderson and Andrew, 1998, Mehler et al., 1997, Haerry et al., 1996, Henderson and Andrew, 1997, Dean, 1996, Hogan, 1996, Kirilly et al., 2006, Eaton and Davis, 2005, Terriente and de Celis, 2007, Kaneko et al., 2011, Serpe and O'Connor, 2006, Xia et al., 2012, Ulvklo et al., 2008, Wang et al., 2011, Nègre et al., 2011, Debat et al., 2009, Vlachos and Harden, 2011, Umemori et al., 2009, Corrigall et al., 2007, Jiang et al., 2008, Tyler and Baker, 2007, Ng, 2008, Goold and Davis, 2007, O'Connor-Giles et al., 2008, Zeng et al., 2008, Xie et al., 2005, Widmann and Dahmann, 2009, Twombly et al., 2009, Ninov et al., 2010, Vuilleumier et al., 2010, Terriente-Félix et al., 2010, Kim et al., 2010, The modENCODE Consortium, 2010, The modENCODE Consortium, 2010, Dworkin et al., 2011, Hill, 1996, Fischer et al., 2012, Seppa et al., 2008, Giorgianni and Mann, 2011, Reddy and Irvine, 2011, Peterson et al., 2012, Chen et al., 2012) MAD (Kockel et al., 2001, Bassett et al., 1997, Chen et al., 2006, Collins et al., 2006, Fuller and Spradling, 2007, Cheesman et al., 2004, Bickel et al., 2008, Kang and Bier, 2010, Szuperák et al., 2011, Gesualdi and Haerry, 2007, Chung et al., 2011, Shravage et al., 2007, Rogulja et al., 2008, Yakoby et al., 2008, Nagaraj and Banerjee, 2009, Niepielko et al., 2011, Mulligan et al., 2012, Haerry, 2010, Iovino et al., 2009, Potthoff and Olson, 2007, MacArthur et al., 2009, Ratnaparkhi et al., 2008, Chen and Xu, 2010, Goldstein et al., 2011, Khuong et al., 2010, Mathur et al., 2010, Rusten et al., 2006, Niepielko et al., 2012) Mat (Koh et al., 2004) pMad (Hatfield et al., 2005, Wang and Ferguson, 2005, Baumgardt et al., 2009, Vrailas and Moses, 2006, Boylan et al., 2008, Yan et al., 2009, Guo and Wang, 2009, Nahmad and Stathopoulos, 2009, Hatton-Ellis et al., 2007, Umulis et al., 2010, Sun et al., 2010, Sopory et al., 2010, Junion et al., 2012, Lu et al., 2012) p-Mad (Adachi-Yamada et al., 2005) pMAD (Kato et al., 2004, Belenkaya et al., 2004, De Graeve et al., 2004, Ward et al., 2006, Bonds et al., 2007) P-mad (López-Onieva et al., 2007) P-Mad (Sotillos and De Celis, 2005) PMad (Wang et al., 2007) pSmad (Leatherman and Dinardo, 2010) Smad (Leatherman and Dinardo, 2010) Smad1 (Chen and Xu, 2010)
Name Synonym	apang   Mother against decapentaplegic (Manjon et al., 2007) mothers against decapentaplegi (Koh et al., 2004) Mothers against Decapentaplegic (Burnett et al., 2006, Chen et al., 2006, Xu et al., 2007) Mothers against decapentaplegic (Vrailas and Moses, 2006, Chen et al., 2010) mothers against decapentaplegic (Lim et al., 2008, Mathur et al., 2010) Mothers Against Decapentaplegic (Higashi-Kovtun et al., 2010, Khuong et al., 2010) Mothers against dpp (Ball et al., 2010, Rawson et al., 2003, Yamashita et al., 2005, Marques, 2005, Bernardi et al., 2006, Philippakis et al., 2006, Bonds et al., 2007, Boylan et al., 2008, Ogiso et al., 2011, Chang et al., 2008, Umemori et al., 2007, Twombly et al., 2009, Stefancsik and Sarkar, 2003, Salzer et al., 2010, Losada-Pérez et al., 2010, Liu et al., 2011, Gao and Laughon, 2007, Aboukhalil and Bulyk, 2012) Mothers against Dpp (Horabin, 2005, Dworkin and Gibson, 2006, Anderson et al., 2006, Letizia et al., 2007, Umemori et al., 2009, Pinto et al., 2008, Evans et al., 2009, Zhao et al., 2008, Weiss et al., 2010, Agelopoulos et al., 2012) Mothers against DPP (Bickel et al., 2008, Lembong et al., 2009) mothers against dpp (Arbouzova and Zeidler, 2006, Corrigall et al., 2007, Firth et al., 2010, Terriente-Félix et al., 2010) Mothers Against Dpp (Wang and Ward, 2010, Rusten et al., 2006) phosphorylated Smad (Baumgardt et al., 2009) phosphoSmad (Leatherman and Dinardo, 2010) phospho-Smad (Leatherman and Dinardo, 2010)
Secondary FlyBase IDs
	FBgn0000105 FBgn0013945 FBgn0022270 FBgn0024356
References ( 736 )
Generate a list of	All references relating to this gene ( 736 )
List References by type	Research paper  ( 443 ) Supplementary material  ( 18 ) Review  ( 97 ) Personal communication to FlyBase  ( 25 ) Abstract  ( 142 ) FlyBase analysis  ( 3 ) Letter  ( 1 ) Protein sequence record  ( 1 ) Website  ( 1 ) Curated genome annotation  ( 1 ) Book  ( 1 ) Conference report  ( 2 ) Thesis  ( 1 )
Recent research papers ( 69 )
	Ables and Drummond-Barbosa, 2013, Development 140(3): 530--540 Cyclin E controls Drosophila female germline stem cell maintenance independently of its role in proliferation by modulating responsiveness to niche signals. [FBrf0220400] Baker, 2013, PLoS ONE 8(3): e58266 Developmental Regulation of Nucleolus Size during Drosophila Eye Differentiation. [FBrf0220989] Gorostiza and Ceriani, 2013, J. Neurosci. 33(2): 687--696 Retrograde bone morphogenetic protein signaling shapes a key circadian pacemaker circuit. [FBrf0220479] Peterson and O'Connor, 2013, Development 140(3): 649--659 Activin receptor inhibition by Smad2 regulates Drosophila wing disc patterning through BMP-response elements. [FBrf0220378] Rockel et al., 2013, Nucleic Acids Res. 41(4): e52 iSLIM: a comprehensive approach to mapping and characterizing gene regulatory networks. [FBrf0220859] Aboukhalil and Bulyk, 2012, Bioinformatics 28(11): 1446--1454 LOESS correction for length variation in gene set-based genomic sequence analysis. [FBrf0218376] Agelopoulos et al., 2012, Cell Rep. 1(4): 350--359 Developmental regulation of chromatin conformation by Hox proteins in Drosophila. [FBrf0218139] Akiyama et al., 2012, Sci. Signal. 5(218): ra28 A Large Bioactive BMP Ligand with Distinct Signaling Properties Is Produced by Alternative Proconvertase Processing. [FBrf0217914] Beck et al., 2012, J. Neurosci. 32(20): 7058--7073 Regulation of Fasciclin II and Synaptic Terminal Development by the Splicing Factor Beag. [FBrf0218385] Cash and Andrews, 2012, BMC Dev. Biol. 12: 4 Fine scale analysis of gene expression in Drosophila melanogaster gonads reveals Programmed cell death 4 promotes the differentiation of female germline stem cells. [FBrf0217547] Chen et al., 2012, Development 139(12): 2170--2176 Crossveinless d is a vitellogenin-like lipoprotein that binds BMPs and HSPGs, and is required for normal BMP signaling in the Drosophila wing. [FBrf0218386] Dahal et al., 2012, Development 139(19): 3653--3664 An inwardly rectifying K+ channel is required for patterning. [FBrf0219361] Eade et al., 2012, PLoS Genet. 8(2): e1002501 Developmental transcriptional networks are required to maintain neuronal subtype identity in the mature nervous system. [FBrf0217668] Fischer et al., 2012, PLoS ONE 7(8): e42349 fussel (fuss) - A Negative Regulator of BMP Signaling in Drosophila melanogaster. [FBrf0219160] Gomez et al., 2012, J. Cell Biol. 199(7): 1131--1143 Tao controls epithelial morphogenesis by promoting Fasciclin 2 endocytosis. [FBrf0220436] Junion et al., 2012, Cell 148(3): 473--486 A transcription factor collective defines cardiac cell fate and reflects lineage history. [FBrf0217404] Kagey et al., 2012, Mech. Dev. 129(9-12): 339--349 Regulation of Yorkie activity in Drosophila imaginal discs by the Hedgehog receptor gene patched. [FBrf0219794] Karim et al., 2012, J. R. Soc. Interface 9(70): 1073--1083 Secreted, receptor-associated bone morphogenetic protein regulators reduce stochastic noise intrinsic to many extracellular morphogen distributions. [FBrf0217860] Kim and Marqués, 2012, Dev. Neurobiol. 72(12): 1541--1558 The Ly6 neurotoxin-like molecule target of wit regulates spontaneous neurotransmitter release at the developing neuromuscular junction in Drosophila. [FBrf0219871] Le and Wharton, 2012, Dev. Dyn. 241(1): 200--214 Hyperactive BMP signaling induced by ALK2(R206H) requires type II receptor function in a Drosophila model for classic fibrodysplasia ossificans progressiva. [FBrf0217004] Lu et al., 2012, PLoS Biol. 10(7): e1001357 Niche-associated activation of rac promotes the asymmetric division of Drosophila female germline stem cells. [FBrf0218889] Mulligan et al., 2012, Proc. Natl. Acad. Sci. U.S.A. 109(2): 370--377 Secreted Wingless-interacting molecule (Swim) promotes long-range signaling by maintaining Wingless solubility. [FBrf0217131] Niepielko et al., 2012, Biophys. J. 102(8): 1722--1730 Evolution of BMP Signaling in Drosophila Oogenesis: A Receptor-Based Mechanism. [FBrf0218807] Pennetier et al., 2012, Proc. Natl. Acad. Sci. U.S.A. 109(9): 3389--3394 Size control of the Drosophila hematopoietic niche by bone morphogenetic protein signaling reveals parallels with mammals. [FBrf0217568] Peterson et al., 2012, PLoS ONE 7(5): e36548 R-smad competition controls activin receptor output in Drosophila. [FBrf0218205] Stinchfield et al., 2012, Development 139(15): 2721--2729 Fat facets deubiquitylation of Medea/Smad4 modulates interpretation of a Dpp morphogen gradient. [FBrf0219593] Sun et al., 2012, PLoS Genet. 8(2): e1002515 Neurophysiological Defects and Neuronal Gene Deregulation in Drosophila mir-124 Mutants. [FBrf0217508] Takaesu et al., 2012, Development 139(18): 3392--3401 Drosophila CORL is required for Smad2-mediated activation of Ecdysone Receptor expression in the mushroom body. [FBrf0219224] Vied et al., 2012, Dev. Cell 23(4): 836--848 Regulation of stem cells by intersecting gradients of long-range niche signals. [FBrf0219709] Xia et al., 2012, Curr. Biol. 22(6): 515--521 The Niche-Dependent Feedback Loop Generates a BMP Activity Gradient to Determine the Germline Stem Cell Fate. [FBrf0217877] Zhu et al., 2012, Development 139(8): 1457--1466 Differential regulation of mesodermal gene expression by Drosophila cell type-specific Forkhead transcription factors. [FBrf0217779] Abruzzi et al., 2011, Genes Dev. 25(22): 2374--2386 Drosophila CLOCK target gene characterization: implications for circadian tissue-specific gene expression. [FBrf0216798] Ben-Zvi et al., 2011, Curr. Biol. 21(16): 1391--1396 Expansion-repression mechanism for scaling the dpp activation gradient in Drosophila wing imaginal discs. [FBrf0214705] Bhattacharya and Baker, 2011, Cell 147(4): 881--892 A Network of Broadly Expressed HLH Genes Regulates Tissue-Specific Cell Fates. [FBrf0216641] Chung et al., 2011, Nat. Methods 8(2): 171--176 A microfluidic array for large-scale ordering and orientation of embryos. [FBrf0212871] Clark et al., 2011, Curr. Biol. 21(19): 1672--1677 Multiple TGF-β Superfamily Signals Modulate the Adult Drosophila Immune Response. [FBrf0216429] Colozza et al., 2011, Tissue Cell 43(4): 254--261 Drosophila nucleoporin Nup154 controls cell viability, proliferation and nuclear accumulation of Mad transcription factor. [FBrf0214248] Dejima et al., 2011, J. Biol. Chem. 286(19): 17103--17111 Novel Contact-dependent Bone Morphogenetic Protein (BMP) Signaling Mediated by Heparan Sulfate Proteoglycans. [FBrf0213647] Dworkin et al., 2011, Genetics 187(4): 1171--1184 The effects of weak genetic perturbations on the transcriptome of the wing imaginal disc and its association with wing shape in Drosophila melanogaster. [FBrf0214376] Eivers et al., 2011, Sci. Signal. 4(194): ra68 Phosphorylation of Mad Controls Competition Between Wingless and BMP Signaling. [FBrf0216411] Gancz et al., 2011, PLoS Biol. 9(11): e1001202 Coordinated regulation of niche and stem cell precursors by hormonal signaling. [FBrf0217934] Giorgianni and Mann, 2011, Dev. Cell 20(4): 455--468 Establishment of Medial Fates along the Proximodistal Axis of the Drosophila Leg through Direct Activation of dachshund by Distalless. [FBrf0213490] Goldstein et al., 2011, Hum. Mol. Genet. 20(5): 894--904 SMAD signaling drives heart and muscle dysfunction in a Drosophila model of muscular dystrophy. [FBrf0212934] Hamaratoglu et al., 2011, PLoS Biol. 9(10): e1001182 Dpp signaling activity requires pentagone to scale with tissue size in the growing Drosophila wing imaginal disc. [FBrf0216502] Harris et al., 2011, Dev. Cell 20(1): 72--83 Brat Promotes Stem Cell Differentiation via Control of a Bistable Switch that Restricts BMP Signaling. [FBrf0212787] Huang et al., 2011, Development 138(11): 2283--2291 DPP-mediated TGF{beta} signaling regulates juvenile hormone biosynthesis by activating the expression of juvenile hormone acid methyltransferase. [FBrf0213687] James and Broihier, 2011, Development 138(15): 3273--3286 Crimpy inhibits the BMP homolog Gbb in motoneurons to enable proper growth control at the Drosophila neuromuscular junction. [FBrf0214390] Kaneko et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(27): 11127--11132 Smad inhibition by the Ste20 kinase Misshapen. [FBrf0214278] König et al., 2011, EMBO J. 30(8): 1549--1562 Ecdysteroids affect Drosophila ovarian stem cell niche formation and early germline differentiation. [FBrf0213531] Layalle et al., 2011, Development 138(11): 2315--2323 Engrailed homeoprotein acts as a signaling molecule in the developing fly. [FBrf0213689] Li et al., 2011, Genome Biol. 12(4): R34 The role of chromatin accessibility in directing the widespread, overlapping patterns of Drosophila transcription factor binding. [FBrf0216471] Liu et al., 2011, Dev. Growth Differ. 53(6): 822--841 Negative modulation of bone morphogenetic protein signaling by Dullard during wing vein formation in Drosophila. [FBrf0214664] Mikhaylova and Nurminsky, 2011, BMC Biol. 9: 29 Lack of global meiotic sex chromosome inactivation, and paucity of tissue-specific gene expression on the Drosophila X chromosome. [FBrf0213839] Miller et al., 2011, J. Neurosci. 31(14): 5335--5347 Drosophila mmp2 regulates the matrix molecule faulty attraction (frac) to promote motor axon targeting in Drosophila. [FBrf0213409] Niepielko et al., 2011, Dev. Biol. 354(1): 151--159 BMP signaling dynamics in the follicle cells of multiple Drosophila species. [FBrf0213633] Nègre et al., 2011, Nature 471(7339): 527--531 A cis-regulatory map of the Drosophila genome. [FBrf0213303] O'Keefe et al., 2011, Mech. Dev. 128(1-2): 59--70 EndoGI modulates Notch signaling and axon guidance in Drosophila. [FBrf0213012] Ogiso et al., 2011, Dev. Growth Differ. 53(5): 668--678 Robustness of the Dpp morphogen activity gradient depends on negative feedback regulation by the inhibitory Smad, Dad. [FBrf0213936] Oh and Irvine, 2011, Dev. Cell 20(1): 109--122 Cooperative Regulation of Growth by Yorkie and Mad through bantam. [FBrf0212763] Peluso et al., 2011, Dev. Cell 21(2): 375--383 Shaping BMP Morphogen Gradients through Enzyme-Substrate Interactions. [FBrf0214661] Qian et al., 2011, Genomics 97(5): 294--303 The full-length transcripts and promoter analysis of intergenic microRNAs in Drosophila melanogaster. [FBrf0213652] Quijano et al., 2011, Genetics 189(3): 809--824 Wg Signaling via Zw3 and Mad Restricts Self-Renewal of Sensory Organ Precursor Cells in Drosophila. [FBrf0216675] Reddy and Irvine, 2011, Development 138(23): 5201--5212 Regulation of Drosophila glial cell proliferation by Merlin-Hippo signaling. [FBrf0216584] Rodal et al., 2011, J. Cell Biol. 193(1): 201--217 A presynaptic endosomal trafficking pathway controls synaptic growth signaling. [FBrf0213353] Rodriguez, 2011, PLoS ONE 6(4): e18418 Drosophila TIEG Is a Modulator of Different Signalling Pathways Involved in Wing Patterning and Cell Proliferation. [FBrf0213463] Szuperák et al., 2011, Development 138(4): 715--724 Feedback regulation of Drosophila BMP signaling by the novel extracellular protein Larval Translucida. [FBrf0212881] Vlachos and Harden, 2011, Genetics 187(2): 501--512 Genetic Evidence for Antagonism Between Pak Protein Kinase and Rho1 Small GTPase Signaling in Regulation of the Actin Cytoskeleton During Drosophila Oogenesis. [FBrf0212983] Wang et al., 2011, Cell Res. 21(4): 700--703 Effective gene silencing in Drosophila ovarian germline by artificial microRNAs. [FBrf0213407] Yuva-Aydemir et al., 2011, J. Neurosci. 31(19): 7005--7015 Spinster Controls Dpp Signaling during Glial Migration in the Drosophila Eye. [FBrf0213705] Show all research papers ...
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