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Gene Dmel\Top2

General Information
Symbol	Dmel\Top2	Species	D. melanogaster
Name	Topoisomerase 2	Annotation symbol	CG10223
Feature type	protein_coding_gene	FlyBase ID	FBgn0003732
Gene Model Status	Current	Stock availability	9 publicly available
Also Known As	topo II, TopoII, CHRAC, dCHRAC, TopII
Genomic Location
Chromosome (arm)	2L	Recombination map
Cytogenetic map	37E1-37E1	Sequence location	2L:19,447,365..19,453,490 [-]
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 Topoisomerase 2 is referred to in FlyBase by the symbol Dmel\Top2 (CG10223, FBgn0003732). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: DNA topoisomerase type II (ATP-hydrolyzing) activity; satellite DNA binding; mRNA binding; rDNA binding; four-way junction DNA binding; DNA binding. There is experimental evidence that it is involved in the biological process: mitotic chromosome condensation; metaphase plate congression; mitotic G2 DNA damage checkpoint; DNA topological change; mitosis; mitotic spindle organization; sister chromatid segregation; chromatin silencing. 28 alleles are reported. The phenotypes of these alleles are annotated with: polytene chromosome; ovary; mesothoracic tergum; female germline stem cell. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: Arginine repressor C-terminal-like domain; DNA topoisomerase II, eukaryotic-type; DNA topoisomerase, type IIA; DNA topoisomerase, type IIA, central domain; DNA topoisomerase, type IIA, conserved site; DNA topoisomerase, type IIA, subunit A/ C-terminal, alpha-beta; DNA topoisomerase, type IIA, subunit A/C-terminal; DNA topoisomerase, type IIA, subunit B, domain 2; DNA topoisomerase, type IIA-like domain; Histidine kinase-like ATPase, ATP-binding domain; Ribosomal protein S5 domain 2-type fold; Ribosomal protein S5 domain 2-type fold, subgroup; Toprim domain; Type IIA DNA topoisomerase subunit A, alpha-helical domain. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of very high expression to a trough of moderate expression. Peak expression observed within 00-12 hour embryonic stages. Summary of FlyAtlas Anatomical Expression Data: Many larval and adult organs/tissues expressed at moderate levels. Expression at high levels in the following post-embryonic organs or tissues: larval central nervous system, larval salivary gland, larval trachea, adult ovary. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, adult eye, adult central nervous system, adult crop, larval/adult midgut, larval/adult hindgut, larval/adult Malpighian tubules, larval fat body, adult salivary gland, adult spermathecae, adult male reproductive system, larval carcass. Comments on Affy2 ProbeSet: ProbeSet 1631837_at completely aligns to an exonic region of the only FlyBase-annotated transcript isoform of Top2. Gene sequence location is 2L:19447365..19453490.
User Contributed Data
	Beta version, feedback welcome
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	Top2: Topoisomerase 2 Top2 is an essential gene that encodes the large subunit of type II DNA topoisomerase, an enzyme believed to play an important role in the condensation, decondensation, and segregation of chromosomes. The enzyme is a major component of the nuclear matrix of Drosophila cells (Berrios et al., 1985) and is distributed along polytene chromosomes paralleling the distribution of the DNA (Heller et al., 1986). It is believed to act by passing a DNA segment through a transient double-stranded break in another segment. Major cleavage sites for type II topisomerase have been found in nontranscribed spacer segments and in the 5' and 3' ends of Hsp70 and the histone genes (Udvardy et al., 1985). When prepared from embryos, the purified enzyme is made up of a major polypeptide encoded by Top2 of 166,000 daltons, with binding sites for both DNA and ATP, and, in addition, smaller polypeptides of 30,000-40,000 and 132,000-145,000 daltons (Sander and Hsieh, 1983; Shelton et al., 1983; Heller et al., 1986). Protein kinase activity is associated with Drosophila topoisomerase II (Sander et al., 1984; Ackerman, Glover, and Osheroff, 1985, Proc. Nat. Acad. Sci. USA 82: 3164-68).
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
FB2013_02	Controlled Vocabulary Terms mitotic cell cycle G2/M transition decatenation checkpoint resolution of meiotic recombination intermediates mitotic recombination DNA-dependent DNA replication nucleoid synaptonemal complex
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 37E1-37E1   Limits computationally determined from genome sequence between P{EP}dntEP2158 and P{EP}EP623&P{PZ}spi01068
Experimentally Determined Cytological Location
Cytogenetic map Notes References 37D2-37D6   (determined by in situ hybridisation)   (Wyckoff et al., 1989) 37D2-37D6   (determined by in situ hybridisation)   (Hsieh et al., 1987) 37D2-37D6   (determined by in situ hybridisation)   (Nolan et al., 1986)
Experimentally Determined Recombination Data
Location
Left of (cM)
Right of (cM)
Notes	Maps very close to RanGap.
Gene Model & Products
Please see the GBrowse view of Dmel\Top2 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 Annotated transcripts do not represent all supported alternative splices within 5' UTR.
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) Top2-RA FBtr0081287  NM_057412   4935   1447 Top2-RB FBtr0332320  NM_001273651   5242   1447
Additional Transcript Data & Comments
Reported size (kB)	5.1 (northern blot) (Nolan et al., 1986)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein Top2-PA   FBpp0080825   164.4   1447   8.45     NP_476760   AAF53802 Top2-PB   FBpp0304598   164.4   1447   8.45     NP_001260580   AGB93115
Additional Polypeptide Data & Comments
Reported size (kDa)	1447 (aa); 164.424 (kD predicted) (Wyckoff et al., 1989) 170 (kD predicted) (Nolan et al., 1986)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • DNA topoisomerase, type IIA, conserved site (IPR018522) Toprim domain (IPR006171) DNA topoisomerase II, eukaryotic-type (IPR001154) DNA topoisomerase, type IIA (IPR001241) DNA topoisomerase, type IIA, subunit A/C-terminal (IPR002205) Histidine kinase-like ATPase, ATP-binding domain (IPR003594) DNA topoisomerase, type IIA, subunit B, domain 2 (IPR013506) DNA topoisomerase, type IIA, subunit A/ C-terminal, alpha-beta (IPR013758) DNA topoisomerase, type IIA, central domain (IPR013759) DNA topoisomerase, type IIA-like domain (IPR013760) Type IIA DNA topoisomerase subunit A, alpha-helical domain (IPR013757) Ribosomal protein S5 domain 2-type fold (IPR020568) Arginine repressor C-terminal-like domain (IPR024946) Ribosomal protein S5 domain 2-type fold, subgroup (IPR014721)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA201586   AA202773   AA246220   AA246718   AA264428   AA265000   AA390349   AA390370   AA392310   AA392354   AA392649   AA392776   AA438611   AA440367   AA440850   AA536447   AA539227   AA539836   AA540137   AA540322   AA540836   AA567177   AA695700   AA696280   AA735593   AA803107   AA804038   AA816358   AA817201   AA817550   AA817557   AA820814   AA942028   AA942335   AA950010   AA951624   AA952140   AA990856   AE014134 AAF53802   AC005428   AC007082   AE014134 AGB93115   AI062052   AI113517   AI113527   AI134063   AI456060   AI512878   AI515012   AI518809   AI519483   AI543820   AI546459   AX093881   BG633850   BG638225   BG638415   BG638536   BI166733   BI215979   BI229518   BI232953   BI234659   BI236713   BI237771   BI354258   BI354331   BI354342   BI358681   BI364637   BI364649   BI373398   BP543983   BP543984   BP554184   BP558092   BP558093   BQ103413   BT010240 AAQ23558   CK132442   CO178066   CO178265   CO190687   CO264422   CO273989   CO286780   CO297807   CO298690   CO300949   CO308231   CO312678   CO314551   CO323924   CO324900   CZ469933   CZ474486   CZ487766   DU708674   EC053114   EC053608   EC054092   EC056487   EC057704   EC057712   EC058298   EC060568   EC063127   EC064348   EC068665   EC076463   EC077634   EC080909   EC199996   EC209612   EC212093   EC218340   EC221225   EC244322   EC246805   EC248404   EC262527   EV581208   FE862619   FE862637   FE862702   FE862721   FE862742   FE862751   FE863183   FE863211   FE863279   FE863325   GH739303   GH739913   GH740921   GH743794   GH743893   GH744519   GH744762   GH746914   GH761514   GH762047   GH762161   GH762361   GH762519   GH764641   GH768097   GH769712   GO254513   GO254533   U53787   U53788   U53789   U53790   U53791   U53792   U53793   U53794   U53795   U53796   U53797   U53798   X61209 CAA43523
UniProtKB/Swiss-Prot	P15348
UniProtKB/TrEMBL	Q6NR40
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
Additional Descriptive Data
Marker for
Subcellular Localization
CV Term
Polypeptide Expression
mass spectroscopy Stage Tissue/Position (including subcellular localization) Reference day 7 of adulthood -- day 49 of adulthood   adult heart   (Cammarato et al., 2011)
Additional Descriptive Data
Marker for
Subcellular Localization (GO Cellular Component)
CV term Evidence References
NOT ACF complex inferred from direct assay (Ito et al., 1997) CHRAC inferred from direct assay (Varga-Weisz et al., 1997) NOT CHRAC inferred from direct assay (Eberharter et al., 2001) chromosome inferred from direct assay (Zhang et al., 2000) nucleus inferred from direct assay (Colflesh et al., 1999, Zhang et al., 2000, Lee et al., 1989, Gemkow et al., 2001)
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: Many larval and adult organs/tissues expressed at moderate levels. Expression at high levels in the following post-embryonic organs or tissues: larval central nervous system, larval salivary gland, larval trachea, adult ovary. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, adult eye, adult central nervous system, adult crop, larval/adult midgut, larval/adult hindgut, larval/adult Malpighian tubules, larval fat body, adult salivary gland, adult spermathecae, adult male reproductive system, larval 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    1372.025 Larval Midgut    155.5 Larval Hindgut    226.9 Larval Malpighian Tubules    306.6 Larval Fat Body    112 Larval Salivary Gland    517.9 Larval Trachea    593.35 Larval Carcass    296.175 Adult Head    125.4 Adult Eye    151.275 Adult Brain    219.1 Adult Thoracic-Abdominal Ganglion    219.6 Adult Crop    137 Adult Midgut    105.6 Adult Hindgut    126.2 Adult Malpighian Tubules    166.5 Adult Fat Body    77.7 Adult Salivary Gland    144.1 Adult Heart    97.6 Adult VirginFemale Spermatheca    189.9 Adult InseminatedFemale Spermatheca    217.3 Adult Ovary    1042.3 Adult Testis    196 Adult Male Accessory Gland    373.4 Adult Carcass    99.6 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System  (1372.025) Larval Midgut    155.5 Larval Hindgut    226.9 Larval Malpighian Tubules    306.6 Larval Fat Body    112 Larval Salivary Gland  517.9 Larval Trachea  (593.35) Larval Carcass    296.175 Adult Head    125.4 Adult Eye    151.275 Adult Brain    219.1 Adult Thoracic-Abdominal Ganglion    219.6 Adult Crop    137 Adult Midgut    105.6 Adult Hindgut    126.2 Adult Malpighian Tubules    166.5 Adult Fat Body    77.7 Adult Salivary Gland    144.1 Adult Heart    97.6 Adult VirginFemale Spermatheca    189.9 Adult InseminatedFemale Spermatheca    217.3 Adult Ovary  (1042.3) Adult Testis    196 Adult Male Accessory Gland    373.4 Adult Carcass    99.6 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System  (1372.025) Larval Midgut    155.5 Larval Hindgut    226.9 Larval Malpighian Tubules    306.6 Larval Fat Body    112 Larval Salivary Gland    517.9 Larval Trachea    593.35 Larval Carcass    296.175 Adult Head    125.4 Adult Eye    151.275 Adult Brain    219.1 Adult Thoracic-Abdominal Ganglion    219.6 Adult Crop    137 Adult Midgut    105.6 Adult Hindgut    126.2 Adult Malpighian Tubules    166.5 Adult Fat Body    77.7 Adult Salivary Gland    144.1 Adult Heart    97.6 Adult VirginFemale Spermatheca    189.9 Adult InseminatedFemale Spermatheca    217.3 Adult Ovary  1042.3 Adult Testis    196 Adult Male Accessory Gland    373.4 Adult Carcass    99.6 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    1372.025 Larval Midgut    155.5 Larval Hindgut    226.9 Larval Malpighian Tubules    306.6 Larval Fat Body    112 Larval Salivary Gland    517.9 Larval Trachea    593.35 Larval Carcass    296.175 Adult Head    125.4 Adult Eye    151.275 Adult Brain    219.1 Adult Thoracic-Abdominal Ganglion    219.6 Adult Crop    137 Adult Midgut    105.6 Adult Hindgut    126.2 Adult Malpighian Tubules    166.5 Adult Fat Body    77.7 Adult Salivary Gland    144.1 Adult Heart    97.6 Adult VirginFemale Spermatheca    189.9 Adult InseminatedFemale Spermatheca    217.3 Adult Ovary    1042.3 Adult Testis    196 Adult Male Accessory Gland    373.4 Adult Carcass    99.6 Expression Level Scale  Very high  log, scaled to maximum expression level Tissue   Expression Level Larval Central Nervous System    1372.025 Larval Midgut    155.5 Larval Hindgut    226.9 Larval Malpighian Tubules    306.6 Larval Fat Body    112 Larval Salivary Gland    517.9 Larval Trachea    593.35 Larval Carcass    296.175 Adult Head    125.4 Adult Eye    151.275 Adult Brain    219.1 Adult Thoracic-Abdominal Ganglion    219.6 Adult Crop    137 Adult Midgut    105.6 Adult Hindgut    126.2 Adult Malpighian Tubules    166.5 Adult Fat Body    77.7 Adult Salivary Gland    144.1 Adult Heart    97.6 Adult VirginFemale Spermatheca    189.9 Adult InseminatedFemale Spermatheca    217.3 Adult Ovary    1042.3 Adult Testis    196 Adult Male Accessory Gland    373.4 Adult Carcass    99.6 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System  (1372.025) Larval Midgut    155.5 Larval Hindgut    226.9 Larval Malpighian Tubules    306.6 Larval Fat Body    112 Larval Salivary Gland  517.9 Larval Trachea  593.35 Larval Carcass    296.175 Adult Head    125.4 Adult Eye    151.275 Adult Brain    219.1 Adult Thoracic-Abdominal Ganglion    219.6 Adult Crop    137 Adult Midgut    105.6 Adult Hindgut    126.2 Adult Malpighian Tubules    166.5 Adult Fat Body    77.7 Adult Salivary Gland    144.1 Adult Heart    97.6 Adult VirginFemale Spermatheca    189.9 Adult InseminatedFemale Spermatheca    217.3 Adult Ovary  (1042.3) Adult Testis    196 Adult Male Accessory Gland    373.4 Adult Carcass    99.6 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System  1372.025 Larval Midgut    155.5 Larval Hindgut    226.9 Larval Malpighian Tubules    306.6 Larval Fat Body    112 Larval Salivary Gland    517.9 Larval Trachea    593.35 Larval Carcass    296.175 Adult Head    125.4 Adult Eye    151.275 Adult Brain    219.1 Adult Thoracic-Abdominal Ganglion    219.6 Adult Crop    137 Adult Midgut    105.6 Adult Hindgut    126.2 Adult Malpighian Tubules    166.5 Adult Fat Body    77.7 Adult Salivary Gland    144.1 Adult Heart    97.6 Adult VirginFemale Spermatheca    189.9 Adult InseminatedFemale Spermatheca    217.3 Adult Ovary  1042.3 Adult Testis    196 Adult Male Accessory Gland    373.4 Adult Carcass    99.6 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    1372.025 Larval Midgut    155.5 Larval Hindgut    226.9 Larval Malpighian Tubules    306.6 Larval Fat Body    112 Larval Salivary Gland    517.9 Larval Trachea    593.35 Larval Carcass    296.175 Adult Head    125.4 Adult Eye    151.275 Adult Brain    219.1 Adult Thoracic-Abdominal Ganglion    219.6 Adult Crop    137 Adult Midgut    105.6 Adult Hindgut    126.2 Adult Malpighian Tubules    166.5 Adult Fat Body    77.7 Adult Salivary Gland    144.1 Adult Heart    97.6 Adult VirginFemale Spermatheca    189.9 Adult InseminatedFemale Spermatheca    217.3 Adult Ovary    1042.3 Adult Testis    196 Adult Male Accessory Gland    373.4 Adult Carcass    99.6 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    125.4   NA  Eye    151.275   NA  Brain    219.1   1372.025  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    219.6   NA  Crop    137   155.5  Midgut    105.6   226.9  Hindgut    126.2   306.6  Malpighian Tubules    166.5   112  Fat Body    77.7   517.9  Salivary Gland    144.1   NA  Heart    97.6   593.35  Trachea    NA   NA  VirginFemale Spermatheca    189.9   NA  InseminatedFemale Spermatheca    217.3   NA  Ovary    1042.3   NA  Testis    196   NA  Male Accessory Gland    373.4   296.175  Carcass    99.6 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    66 central nervous system, larvae L3    82 central nervous system, pupae P8    21 head, virgin 1-day female    5 head, virgin 4-day female    6 head, virgin 20-day female    5 head, mated 1-day female    5 head, mated 4-day female    6 head, mated 20-day female    5 head, mated 1-day male    6 head, mated 4-day male    8 head, mated 20-day male    11 salivary gland, larvae L3 wandering    1 salivary gland, white prepupae    4 digestive system, larvae L3 wandering    9 digestive system, 1-day adult    4 digestive system, 4-day adult    9 digestive system, 20-day adult    7 fat body, larvae L3 wandering    3 fat body, white prepupae    13 fat body, pupae P8    8 carcass, larvae L3 wandering    49 carcass, 1-day adult    5 carcass, 4-day adult    10 carcass, 20-day adult    7 ovary, virgin 4-day female    156 ovary, mated 4-day female    129 testis, mated 4-day male    11 accessory gland, mated 4-day male    7 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Moderate expression Tissue   Expression Level imaginal disc, larvae L3 wandering  (66) central nervous system, larvae L3  (82) central nervous system, pupae P8    21 head, virgin 1-day female    5 head, virgin 4-day female    6 head, virgin 20-day female    5 head, mated 1-day female    5 head, mated 4-day female    6 head, mated 20-day female    5 head, mated 1-day male    6 head, mated 4-day male    8 head, mated 20-day male    11 salivary gland, larvae L3 wandering    1 salivary gland, white prepupae    4 digestive system, larvae L3 wandering    9 digestive system, 1-day adult    4 digestive system, 4-day adult    9 digestive system, 20-day adult    7 fat body, larvae L3 wandering    3 fat body, white prepupae    13 fat body, pupae P8    8 carcass, larvae L3 wandering  (49) carcass, 1-day adult    5 carcass, 4-day adult    10 carcass, 20-day adult    7 ovary, virgin 4-day female  (156) ovary, mated 4-day female  (129) testis, mated 4-day male    11 accessory gland, mated 4-day male    7 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Tissue   Expression Level imaginal disc, larvae L3 wandering    66 central nervous system, larvae L3    82 central nervous system, pupae P8    21 head, virgin 1-day female    5 head, virgin 4-day female    6 head, virgin 20-day female    5 head, mated 1-day female    5 head, mated 4-day female    6 head, mated 20-day female    5 head, mated 1-day male    6 head, mated 4-day male    8 head, mated 20-day male    11 salivary gland, larvae L3 wandering    1 salivary gland, white prepupae    4 digestive system, larvae L3 wandering    9 digestive system, 1-day adult    4 digestive system, 4-day adult    9 digestive system, 20-day adult    7 fat body, larvae L3 wandering    3 fat body, white prepupae    13 fat body, pupae P8    8 carcass, larvae L3 wandering    49 carcass, 1-day adult    5 carcass, 4-day adult    10 carcass, 20-day adult    7 ovary, virgin 4-day female  (156) ovary, mated 4-day female  (129) testis, mated 4-day male    11 accessory gland, mated 4-day male    7 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    66 central nervous system, larvae L3    82 central nervous system, pupae P8    21 head, virgin 1-day female    5 head, virgin 4-day female    6 head, virgin 20-day female    5 head, mated 1-day female    5 head, mated 4-day female    6 head, mated 20-day female    5 head, mated 1-day male    6 head, mated 4-day male    8 head, mated 20-day male    11 salivary gland, larvae L3 wandering    1 salivary gland, white prepupae    4 digestive system, larvae L3 wandering    9 digestive system, 1-day adult    4 digestive system, 4-day adult    9 digestive system, 20-day adult    7 fat body, larvae L3 wandering    3 fat body, white prepupae    13 fat body, pupae P8    8 carcass, larvae L3 wandering    49 carcass, 1-day adult    5 carcass, 4-day adult    10 carcass, 20-day adult    7 ovary, virgin 4-day female    156 ovary, mated 4-day female    129 testis, mated 4-day male    11 accessory gland, mated 4-day male    7 Expression Level Scale  Extremely high  log, scaled to maximum expression level Tissue   Expression Level imaginal disc, larvae L3 wandering    66 central nervous system, larvae L3    82 central nervous system, pupae P8    21 head, virgin 1-day female    5 head, virgin 4-day female    6 head, virgin 20-day female    5 head, mated 1-day female    5 head, mated 4-day female    6 head, mated 20-day female    5 head, mated 1-day male    6 head, mated 4-day male    8 head, mated 20-day male    11 salivary gland, larvae L3 wandering    1 salivary gland, white prepupae    4 digestive system, larvae L3 wandering    9 digestive system, 1-day adult    4 digestive system, 4-day adult    9 digestive system, 20-day adult    7 fat body, larvae L3 wandering    3 fat body, white prepupae    13 fat body, pupae P8    8 carcass, larvae L3 wandering    49 carcass, 1-day adult    5 carcass, 4-day adult    10 carcass, 20-day adult    7 ovary, virgin 4-day female    156 ovary, mated 4-day female    129 testis, mated 4-day male    11 accessory gland, mated 4-day male    7 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Moderate expression Tissue   Expression Level imaginal disc, larvae L3 wandering  (66) central nervous system, larvae L3  (82) central nervous system, pupae P8    21 head, virgin 1-day female    5 head, virgin 4-day female    6 head, virgin 20-day female    5 head, mated 1-day female    5 head, mated 4-day female    6 head, mated 20-day female    5 head, mated 1-day male    6 head, mated 4-day male    8 head, mated 20-day male    11 salivary gland, larvae L3 wandering    1 salivary gland, white prepupae    4 digestive system, larvae L3 wandering    9 digestive system, 1-day adult    4 digestive system, 4-day adult    9 digestive system, 20-day adult    7 fat body, larvae L3 wandering    3 fat body, white prepupae    13 fat body, pupae P8    8 carcass, larvae L3 wandering  (49) carcass, 1-day adult    5 carcass, 4-day adult    10 carcass, 20-day adult    7 ovary, virgin 4-day female  (156) ovary, mated 4-day female  (129) testis, mated 4-day male    11 accessory gland, mated 4-day male    7 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Tissue   Expression Level imaginal disc, larvae L3 wandering    66 central nervous system, larvae L3    82 central nervous system, pupae P8    21 head, virgin 1-day female    5 head, virgin 4-day female    6 head, virgin 20-day female    5 head, mated 1-day female    5 head, mated 4-day female    6 head, mated 20-day female    5 head, mated 1-day male    6 head, mated 4-day male    8 head, mated 20-day male    11 salivary gland, larvae L3 wandering    1 salivary gland, white prepupae    4 digestive system, larvae L3 wandering    9 digestive system, 1-day adult    4 digestive system, 4-day adult    9 digestive system, 20-day adult    7 fat body, larvae L3 wandering    3 fat body, white prepupae    13 fat body, pupae P8    8 carcass, larvae L3 wandering    49 carcass, 1-day adult    5 carcass, 4-day adult    10 carcass, 20-day adult    7 ovary, virgin 4-day female  (156) ovary, mated 4-day female  129 testis, mated 4-day male    11 accessory gland, mated 4-day male    7 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    66 central nervous system, larvae L3    82 central nervous system, pupae P8    21 head, virgin 1-day female    5 head, virgin 4-day female    6 head, virgin 20-day female    5 head, mated 1-day female    5 head, mated 4-day female    6 head, mated 20-day female    5 head, mated 1-day male    6 head, mated 4-day male    8 head, mated 20-day male    11 salivary gland, larvae L3 wandering    1 salivary gland, white prepupae    4 digestive system, larvae L3 wandering    9 digestive system, 1-day adult    4 digestive system, 4-day adult    9 digestive system, 20-day adult    7 fat body, larvae L3 wandering    3 fat body, white prepupae    13 fat body, pupae P8    8 carcass, larvae L3 wandering    49 carcass, 1-day adult    5 carcass, 4-day adult    10 carcass, 20-day adult    7 ovary, virgin 4-day female    156 ovary, mated 4-day female    129 testis, mated 4-day male    11 accessory gland, mated 4-day male    7 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 very high expression to a trough of moderate expression. Peak expression observed within 00-12 hour embryonic stages. [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    142 embryo 02-04hr    258 embryo 04-06hr    334 embryo 06-08hr    229 embryo 08-10hr    146 embryo 10-12hr    54 embryo 12-14hr    58 embryo 14-16hr    40 embryo 16-18hr    32 embryo 18-20hr    15 embryo 20-22hr    67 embryo 22-24hr    55 larva L1    10 larva L2    18 larva L3 12hr old    15 larva L3 puffstage 1-2    24 larva L3 puffstage 3-6    39 larva L3 puffstage 7-9    45 white prepupae new    43 white prepupae 12hr    45 white prepupae 24hr    53 pupae 2d postWPP    18 pupae 3d postWPP    7 pupae 4d postWPP    7 adult male 01day    8 adult male 05day    11 adult male 30day    10 adult female 01day    56 adult female 05day    114 adult female 30day    106 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (142) embryo 02-04hr  (258) embryo 04-06hr  (334) embryo 06-08hr  (229) embryo 08-10hr  (146) embryo 10-12hr  (54) embryo 12-14hr  (58) embryo 14-16hr  (40) embryo 16-18hr  (32) embryo 18-20hr    15 embryo 20-22hr  (67) embryo 22-24hr  (55) larva L1    10 larva L2    18 larva L3 12hr old    15 larva L3 puffstage 1-2    24 larva L3 puffstage 3-6  (39) larva L3 puffstage 7-9  (45) white prepupae new  (43) white prepupae 12hr  (45) white prepupae 24hr  (53) pupae 2d postWPP    18 pupae 3d postWPP    7 pupae 4d postWPP    7 adult male 01day    8 adult male 05day    11 adult male 30day    10 adult female 01day  (56) adult female 05day  (114) adult female 30day  (106) Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr  (142) embryo 02-04hr  (258) embryo 04-06hr  (334) embryo 06-08hr  (229) embryo 08-10hr  (146) embryo 10-12hr    54 embryo 12-14hr    58 embryo 14-16hr    40 embryo 16-18hr    32 embryo 18-20hr    15 embryo 20-22hr    67 embryo 22-24hr    55 larva L1    10 larva L2    18 larva L3 12hr old    15 larva L3 puffstage 1-2    24 larva L3 puffstage 3-6    39 larva L3 puffstage 7-9    45 white prepupae new    43 white prepupae 12hr    45 white prepupae 24hr    53 pupae 2d postWPP    18 pupae 3d postWPP    7 pupae 4d postWPP    7 adult male 01day    8 adult male 05day    11 adult male 30day    10 adult female 01day    56 adult female 05day  (114) adult female 30day  106 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    142 embryo 02-04hr    258 embryo 04-06hr    334 embryo 06-08hr    229 embryo 08-10hr    146 embryo 10-12hr    54 embryo 12-14hr    58 embryo 14-16hr    40 embryo 16-18hr    32 embryo 18-20hr    15 embryo 20-22hr    67 embryo 22-24hr    55 larva L1    10 larva L2    18 larva L3 12hr old    15 larva L3 puffstage 1-2    24 larva L3 puffstage 3-6    39 larva L3 puffstage 7-9    45 white prepupae new    43 white prepupae 12hr    45 white prepupae 24hr    53 pupae 2d postWPP    18 pupae 3d postWPP    7 pupae 4d postWPP    7 adult male 01day    8 adult male 05day    11 adult male 30day    10 adult female 01day    56 adult female 05day    114 adult female 30day    106 Expression Level Scale  Extremely high  log, scaled to maximum expression level Developmental Stage   Expression Level embryo 00-02hr    142 embryo 02-04hr    258 embryo 04-06hr    334 embryo 06-08hr    229 embryo 08-10hr    146 embryo 10-12hr    54 embryo 12-14hr    58 embryo 14-16hr    40 embryo 16-18hr    32 embryo 18-20hr    15 embryo 20-22hr    67 embryo 22-24hr    55 larva L1    10 larva L2    18 larva L3 12hr old    15 larva L3 puffstage 1-2    24 larva L3 puffstage 3-6    39 larva L3 puffstage 7-9    45 white prepupae new    43 white prepupae 12hr    45 white prepupae 24hr    53 pupae 2d postWPP    18 pupae 3d postWPP    7 pupae 4d postWPP    7 adult male 01day    8 adult male 05day    11 adult male 30day    10 adult female 01day    56 adult female 05day    114 adult female 30day    106 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (142) embryo 02-04hr  (258) embryo 04-06hr  (334) embryo 06-08hr  (229) embryo 08-10hr  (146) embryo 10-12hr  (54) embryo 12-14hr  (58) embryo 14-16hr  (40) embryo 16-18hr  32 embryo 18-20hr    15 embryo 20-22hr  (67) embryo 22-24hr  (55) larva L1    10 larva L2    18 larva L3 12hr old    15 larva L3 puffstage 1-2    24 larva L3 puffstage 3-6  (39) larva L3 puffstage 7-9  (45) white prepupae new  (43) white prepupae 12hr  (45) white prepupae 24hr  (53) pupae 2d postWPP    18 pupae 3d postWPP    7 pupae 4d postWPP    7 adult male 01day    8 adult male 05day    11 adult male 30day    10 adult female 01day  (56) adult female 05day  (114) adult female 30day  (106) Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr  142 embryo 02-04hr  (258) embryo 04-06hr  (334) embryo 06-08hr  (229) embryo 08-10hr  (146) embryo 10-12hr    54 embryo 12-14hr    58 embryo 14-16hr    40 embryo 16-18hr    32 embryo 18-20hr    15 embryo 20-22hr    67 embryo 22-24hr    55 larva L1    10 larva L2    18 larva L3 12hr old    15 larva L3 puffstage 1-2    24 larva L3 puffstage 3-6    39 larva L3 puffstage 7-9    45 white prepupae new    43 white prepupae 12hr    45 white prepupae 24hr    53 pupae 2d postWPP    18 pupae 3d postWPP    7 pupae 4d postWPP    7 adult male 01day    8 adult male 05day    11 adult male 30day    10 adult female 01day    56 adult female 05day  114 adult female 30day  106 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    142 embryo 02-04hr    258 embryo 04-06hr    334 embryo 06-08hr    229 embryo 08-10hr    146 embryo 10-12hr    54 embryo 12-14hr    58 embryo 14-16hr    40 embryo 16-18hr    32 embryo 18-20hr    15 embryo 20-22hr    67 embryo 22-24hr    55 larva L1    10 larva L2    18 larva L3 12hr old    15 larva L3 puffstage 1-2    24 larva L3 puffstage 3-6    39 larva L3 puffstage 7-9    45 white prepupae new    43 white prepupae 12hr    45 white prepupae 24hr    53 pupae 2d postWPP    18 pupae 3d postWPP    7 pupae 4d postWPP    7 adult male 01day    8 adult male 05day    11 adult male 30day    10 adult female 01day    56 adult female 05day    114 adult female 30day    106 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+    70 Schneider line 2 Sg4    61 embryonic 1182-4H    65 embryonic GM2    73 embryonic Kc167    39 embryonic S1    95 embryonic S3    110 leg disc CME L1    75 wing disc CME-W2    107 wing disc ML-DmD8    90 wing disc ML-DmD9    51 wing disc ML-DmD16-c3    117 wing disc ML-DmD21    54 wing disc ML-DmD32    49 haltere disc ML-DmD17-c3    99 eye-antennal disc ML-DmD11    90 antennal disc ML-DmD20-c5    52 mixed discs ML-DmD4-c1    68 CNS ML-DmBG1-c1    83 CNS ML-DmBG2-c2    99 tumorous blood cells mbn2    97 ovary fGS/OSS    112 ovary OSC    85 ovary OSS    78 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Moderate expression Cell Line   Expression Level Schneider line 2 S2R+  (70) Schneider line 2 Sg4  (61) embryonic 1182-4H  (65) embryonic GM2  (73) embryonic Kc167  (39) embryonic S1  (95) embryonic S3  (110) leg disc CME L1  (75) wing disc CME-W2  (107) wing disc ML-DmD8  (90) wing disc ML-DmD9  (51) wing disc ML-DmD16-c3  (117) wing disc ML-DmD21  (54) wing disc ML-DmD32  (49) haltere disc ML-DmD17-c3  (99) eye-antennal disc ML-DmD11  (90) antennal disc ML-DmD20-c5  (52) mixed discs ML-DmD4-c1  (68) CNS ML-DmBG1-c1  (83) CNS ML-DmBG2-c2  (99) tumorous blood cells mbn2  (97) ovary fGS/OSS  (112) ovary OSC  (85) ovary OSS  (78) Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Cell Line   Expression Level Schneider line 2 S2R+    70 Schneider line 2 Sg4    61 embryonic 1182-4H    65 embryonic GM2    73 embryonic Kc167    39 embryonic S1    95 embryonic S3  (110) leg disc CME L1    75 wing disc CME-W2  107 wing disc ML-DmD8    90 wing disc ML-DmD9    51 wing disc ML-DmD16-c3  (117) wing disc ML-DmD21    54 wing disc ML-DmD32    49 haltere disc ML-DmD17-c3    99 eye-antennal disc ML-DmD11    90 antennal disc ML-DmD20-c5    52 mixed discs ML-DmD4-c1    68 CNS ML-DmBG1-c1    83 CNS ML-DmBG2-c2    99 tumorous blood cells mbn2    97 ovary fGS/OSS  (112) ovary OSC    85 ovary OSS    78 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+    70 Schneider line 2 Sg4    61 embryonic 1182-4H    65 embryonic GM2    73 embryonic Kc167    39 embryonic S1    95 embryonic S3    110 leg disc CME L1    75 wing disc CME-W2    107 wing disc ML-DmD8    90 wing disc ML-DmD9    51 wing disc ML-DmD16-c3    117 wing disc ML-DmD21    54 wing disc ML-DmD32    49 haltere disc ML-DmD17-c3    99 eye-antennal disc ML-DmD11    90 antennal disc ML-DmD20-c5    52 mixed discs ML-DmD4-c1    68 CNS ML-DmBG1-c1    83 CNS ML-DmBG2-c2    99 tumorous blood cells mbn2    97 ovary fGS/OSS    112 ovary OSC    85 ovary OSS    78 Expression Level Scale  Extremely high  log, scaled to maximum expression level Cell Line   Expression Level Schneider line 2 S2R+    70 Schneider line 2 Sg4    61 embryonic 1182-4H    65 embryonic GM2    73 embryonic Kc167    39 embryonic S1    95 embryonic S3    110 leg disc CME L1    75 wing disc CME-W2    107 wing disc ML-DmD8    90 wing disc ML-DmD9    51 wing disc ML-DmD16-c3    117 wing disc ML-DmD21    54 wing disc ML-DmD32    49 haltere disc ML-DmD17-c3    99 eye-antennal disc ML-DmD11    90 antennal disc ML-DmD20-c5    52 mixed discs ML-DmD4-c1    68 CNS ML-DmBG1-c1    83 CNS ML-DmBG2-c2    99 tumorous blood cells mbn2    97 ovary fGS/OSS    112 ovary OSC    85 ovary OSS    78 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+  (70) Schneider line 2 Sg4  (61) embryonic 1182-4H  (65) embryonic GM2  (73) embryonic Kc167  (39) embryonic S1  (95) embryonic S3  (110) leg disc CME L1  (75) wing disc CME-W2  (107) wing disc ML-DmD8  (90) wing disc ML-DmD9  (51) wing disc ML-DmD16-c3  (117) wing disc ML-DmD21  (54) wing disc ML-DmD32  (49) haltere disc ML-DmD17-c3  (99) eye-antennal disc ML-DmD11  (90) antennal disc ML-DmD20-c5  (52) mixed discs ML-DmD4-c1  (68) CNS ML-DmBG1-c1  (83) CNS ML-DmBG2-c2  (99) tumorous blood cells mbn2  (97) ovary fGS/OSS  (112) ovary OSC  (85) ovary OSS  (78) Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Cell Line   Expression Level Schneider line 2 S2R+    70 Schneider line 2 Sg4    61 embryonic 1182-4H    65 embryonic GM2    73 embryonic Kc167    39 embryonic S1    95 embryonic S3  110 leg disc CME L1    75 wing disc CME-W2  107 wing disc ML-DmD8    90 wing disc ML-DmD9    51 wing disc ML-DmD16-c3  117 wing disc ML-DmD21    54 wing disc ML-DmD32    49 haltere disc ML-DmD17-c3    99 eye-antennal disc ML-DmD11    90 antennal disc ML-DmD20-c5    52 mixed discs ML-DmD4-c1    68 CNS ML-DmBG1-c1    83 CNS ML-DmBG2-c2    99 tumorous blood cells mbn2    97 ovary fGS/OSS  112 ovary OSC    85 ovary OSS    78 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+    70 Schneider line 2 Sg4    61 embryonic 1182-4H    65 embryonic GM2    73 embryonic Kc167    39 embryonic S1    95 embryonic S3    110 leg disc CME L1    75 wing disc CME-W2    107 wing disc ML-DmD8    90 wing disc ML-DmD9    51 wing disc ML-DmD16-c3    117 wing disc ML-DmD21    54 wing disc ML-DmD32    49 haltere disc ML-DmD17-c3    99 eye-antennal disc ML-DmD11    90 antennal disc ML-DmD20-c5    52 mixed discs ML-DmD4-c1    68 CNS ML-DmBG1-c1    83 CNS ML-DmBG2-c2    99 tumorous blood cells mbn2    97 ovary fGS/OSS    112 ovary OSC    85 ovary OSS    78 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    28 cold shock, 4-day adult    22 heat shock, 4-day adult    45 Cadmium 50 mM 6 hrs, larvae L3    12 Cadmium 50 mM 12 hrs, larvae L3    17 Cadmium 50 mM 48 hrs, 4-day adult    35 Cadmium 100 mM 48 hrs, 4-day adult    83 Copper 0.5 mM 12 hrs, larvae L3    11 Copper 15 mM 48 hrs, 4-day adult    57 Zinc 5 mM 12 hrs, larvae L3    13 Zinc 4.5 mM 48 hrs, 4-day adult    23 Ethanol 2.5% 3 hrs, larvae L3    87 Ethanol 5% 3 hrs, larvae L3    15 Ethanol 10% 3 hrs, larvae L3    11 Caffeine 1.5 mg/ml 4 hrs, larvae L3    15 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    69 Caffeine 25 mg/ml 48 hrs, 4-day adult    29 Paraquat 5 mM 48 hrs, 4-day adult    63 Paraquat 10 mM 48 hrs, 4-day adult    53 Rotenone 2 μg 12 hrs, larvae L3    13 Rotenone 8 μg 12 hrs, larvae L3    7 Expression Level Scale  Very low   Low   Moderate   Moderately high   High  Linear, scaled to Moderate expression Treatment   Expression Level extended cold, 4-day adult  (28) cold shock, 4-day adult    22 heat shock, 4-day adult  (45) Cadmium 50 mM 6 hrs, larvae L3    12 Cadmium 50 mM 12 hrs, larvae L3    17 Cadmium 50 mM 48 hrs, 4-day adult  (35) Cadmium 100 mM 48 hrs, 4-day adult  (83) Copper 0.5 mM 12 hrs, larvae L3    11 Copper 15 mM 48 hrs, 4-day adult  (57) Zinc 5 mM 12 hrs, larvae L3    13 Zinc 4.5 mM 48 hrs, 4-day adult    23 Ethanol 2.5% 3 hrs, larvae L3  (87) Ethanol 5% 3 hrs, larvae L3    15 Ethanol 10% 3 hrs, larvae L3    11 Caffeine 1.5 mg/ml 4 hrs, larvae L3    15 Caffeine 2.5 mg/ml 48 hrs, 4-day adult  (69) Caffeine 25 mg/ml 48 hrs, 4-day adult  (29) Paraquat 5 mM 48 hrs, 4-day adult  (63) Paraquat 10 mM 48 hrs, 4-day adult  (53) Rotenone 2 μg 12 hrs, larvae L3    13 Rotenone 8 μg 12 hrs, larvae L3    7 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Treatment   Expression Level extended cold, 4-day adult    28 cold shock, 4-day adult    22 heat shock, 4-day adult    45 Cadmium 50 mM 6 hrs, larvae L3    12 Cadmium 50 mM 12 hrs, larvae L3    17 Cadmium 50 mM 48 hrs, 4-day adult    35 Cadmium 100 mM 48 hrs, 4-day adult    83 Copper 0.5 mM 12 hrs, larvae L3    11 Copper 15 mM 48 hrs, 4-day adult    57 Zinc 5 mM 12 hrs, larvae L3    13 Zinc 4.5 mM 48 hrs, 4-day adult    23 Ethanol 2.5% 3 hrs, larvae L3    87 Ethanol 5% 3 hrs, larvae L3    15 Ethanol 10% 3 hrs, larvae L3    11 Caffeine 1.5 mg/ml 4 hrs, larvae L3    15 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    69 Caffeine 25 mg/ml 48 hrs, 4-day adult    29 Paraquat 5 mM 48 hrs, 4-day adult    63 Paraquat 10 mM 48 hrs, 4-day adult    53 Rotenone 2 μg 12 hrs, larvae L3    13 Rotenone 8 μg 12 hrs, larvae L3    7 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    28 cold shock, 4-day adult    22 heat shock, 4-day adult    45 Cadmium 50 mM 6 hrs, larvae L3    12 Cadmium 50 mM 12 hrs, larvae L3    17 Cadmium 50 mM 48 hrs, 4-day adult    35 Cadmium 100 mM 48 hrs, 4-day adult    83 Copper 0.5 mM 12 hrs, larvae L3    11 Copper 15 mM 48 hrs, 4-day adult    57 Zinc 5 mM 12 hrs, larvae L3    13 Zinc 4.5 mM 48 hrs, 4-day adult    23 Ethanol 2.5% 3 hrs, larvae L3    87 Ethanol 5% 3 hrs, larvae L3    15 Ethanol 10% 3 hrs, larvae L3    11 Caffeine 1.5 mg/ml 4 hrs, larvae L3    15 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    69 Caffeine 25 mg/ml 48 hrs, 4-day adult    29 Paraquat 5 mM 48 hrs, 4-day adult    63 Paraquat 10 mM 48 hrs, 4-day adult    53 Rotenone 2 μg 12 hrs, larvae L3    13 Rotenone 8 μg 12 hrs, larvae L3    7 Expression Level Scale  Extremely high  log, scaled to maximum expression level Treatment   Expression Level extended cold, 4-day adult    28 cold shock, 4-day adult    22 heat shock, 4-day adult    45 Cadmium 50 mM 6 hrs, larvae L3    12 Cadmium 50 mM 12 hrs, larvae L3    17 Cadmium 50 mM 48 hrs, 4-day adult    35 Cadmium 100 mM 48 hrs, 4-day adult    83 Copper 0.5 mM 12 hrs, larvae L3    11 Copper 15 mM 48 hrs, 4-day adult    57 Zinc 5 mM 12 hrs, larvae L3    13 Zinc 4.5 mM 48 hrs, 4-day adult    23 Ethanol 2.5% 3 hrs, larvae L3    87 Ethanol 5% 3 hrs, larvae L3    15 Ethanol 10% 3 hrs, larvae L3    11 Caffeine 1.5 mg/ml 4 hrs, larvae L3    15 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    69 Caffeine 25 mg/ml 48 hrs, 4-day adult    29 Paraquat 5 mM 48 hrs, 4-day adult    63 Paraquat 10 mM 48 hrs, 4-day adult    53 Rotenone 2 μg 12 hrs, larvae L3    13 Rotenone 8 μg 12 hrs, larvae L3    7 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Moderate expression Treatment   Expression Level extended cold, 4-day adult  28 cold shock, 4-day adult    22 heat shock, 4-day adult  (45) Cadmium 50 mM 6 hrs, larvae L3    12 Cadmium 50 mM 12 hrs, larvae L3    17 Cadmium 50 mM 48 hrs, 4-day adult  (35) Cadmium 100 mM 48 hrs, 4-day adult  (83) Copper 0.5 mM 12 hrs, larvae L3    11 Copper 15 mM 48 hrs, 4-day adult  (57) Zinc 5 mM 12 hrs, larvae L3    13 Zinc 4.5 mM 48 hrs, 4-day adult    23 Ethanol 2.5% 3 hrs, larvae L3  (87) Ethanol 5% 3 hrs, larvae L3    15 Ethanol 10% 3 hrs, larvae L3    11 Caffeine 1.5 mg/ml 4 hrs, larvae L3    15 Caffeine 2.5 mg/ml 48 hrs, 4-day adult  (69) Caffeine 25 mg/ml 48 hrs, 4-day adult  29 Paraquat 5 mM 48 hrs, 4-day adult  (63) Paraquat 10 mM 48 hrs, 4-day adult  (53) Rotenone 2 μg 12 hrs, larvae L3    13 Rotenone 8 μg 12 hrs, larvae L3    7 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Treatment   Expression Level extended cold, 4-day adult    28 cold shock, 4-day adult    22 heat shock, 4-day adult    45 Cadmium 50 mM 6 hrs, larvae L3    12 Cadmium 50 mM 12 hrs, larvae L3    17 Cadmium 50 mM 48 hrs, 4-day adult    35 Cadmium 100 mM 48 hrs, 4-day adult    83 Copper 0.5 mM 12 hrs, larvae L3    11 Copper 15 mM 48 hrs, 4-day adult    57 Zinc 5 mM 12 hrs, larvae L3    13 Zinc 4.5 mM 48 hrs, 4-day adult    23 Ethanol 2.5% 3 hrs, larvae L3    87 Ethanol 5% 3 hrs, larvae L3    15 Ethanol 10% 3 hrs, larvae L3    11 Caffeine 1.5 mg/ml 4 hrs, larvae L3    15 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    69 Caffeine 25 mg/ml 48 hrs, 4-day adult    29 Paraquat 5 mM 48 hrs, 4-day adult    63 Paraquat 10 mM 48 hrs, 4-day adult    53 Rotenone 2 μg 12 hrs, larvae L3    13 Rotenone 8 μg 12 hrs, larvae L3    7 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    28 cold shock, 4-day adult    22 heat shock, 4-day adult    45 Cadmium 50 mM 6 hrs, larvae L3    12 Cadmium 50 mM 12 hrs, larvae L3    17 Cadmium 50 mM 48 hrs, 4-day adult    35 Cadmium 100 mM 48 hrs, 4-day adult    83 Copper 0.5 mM 12 hrs, larvae L3    11 Copper 15 mM 48 hrs, 4-day adult    57 Zinc 5 mM 12 hrs, larvae L3    13 Zinc 4.5 mM 48 hrs, 4-day adult    23 Ethanol 2.5% 3 hrs, larvae L3    87 Ethanol 5% 3 hrs, larvae L3    15 Ethanol 10% 3 hrs, larvae L3    11 Caffeine 1.5 mg/ml 4 hrs, larvae L3    15 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    69 Caffeine 25 mg/ml 48 hrs, 4-day adult    29 Paraquat 5 mM 48 hrs, 4-day adult    63 Paraquat 10 mM 48 hrs, 4-day adult    53 Rotenone 2 μg 12 hrs, larvae L3    13 Rotenone 8 μg 12 hrs, larvae L3    7 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	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003732 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG10223-RA
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal (with Df(2L)Exel9043) Top217-1 Top217-3 Top217-5 Top217-6 Top217-7 Top235-1 Top235-2 Top235-3 Top235-5 Top235-6 Top235-12 Top235-13 Top235-14 lethal (with Top217-1) Top235-13 lethal (with Top217-6) Top235-5 lethal (with Top235-5) Top217-6 lethal (with Top235-13) Top217-1 lethal | heat sensitive (with Df(2L)Exel9043) Top217-2 lethal | recessive Top217-1 Top217-3 Top217-6 Top235-1 Top235-5 Top235-13 partially lethal - majority die (with Df(2L)Exel9043) Top235-4 partially lethal - majority die (with Top217-1) Top217-3 Top217-6 Top235-1 partially lethal - majority die (with Top217-3) Top217-1 Top217-6 Top235-1 Top235-13 partially lethal - majority die (with Top217-6) Top217-1 Top217-3 Top235-1 Top235-13 partially lethal - majority die (with Top235-1) Top217-1 Top217-3 Top217-6 Top235-5 Top235-13 partially lethal - majority die (with Top235-5) Top235-1 partially lethal - majority die (with Top235-13) Top217-3 Top217-6 Top235-1 viable Top2LA00892 viable, with Scer\GAL4pnr-MD237 Top2GD4570 Sterility Allele female sterile (with Top217-1) Top235-1 female sterile (with Top217-3) Top217-6 Top235-1 Top235-5 Top235-13 female sterile (with Top217-6) Top217-3 Top235-1 Top235-13 female sterile (with Top235-1) Top217-1 Top217-3 Top217-6 Top235-5 Top235-13 female sterile (with Top235-5) Top217-3 Top235-1 Top235-13 female sterile (with Top235-13) Top217-3 Top217-6 Top235-1 Top235-5 male fertile (with Top217-1) Top217-3 Top217-6 Top235-1 Top235-5 male fertile (with Top217-3) Top217-1 Top217-6 Top235-5 Top235-13 male fertile (with Top217-6) Top217-1 Top217-3 Top235-1 Top235-13 male fertile (with Top235-1) Top217-1 Top217-6 Top235-5 Top235-13 male fertile (with Top235-5) Top217-1 Top217-3 Top235-1 Top235-13 male fertile (with Top235-13) Top217-3 Top217-6 Top235-1 Top235-5 male sterile (with Top217-3) Top235-1 male sterile (with Top235-1) Top217-3 Other Phenotypes Allele neuroanatomy defective | somatic clone Top2unspecified visible, with Scer\GAL4pnr-MD237 Top2GD4570 Phenotype manifest in Allele embryonic/larval brain | third instar larval stage | heat sensitive Top217-2 female germline stem cell (with Top217-6) Top235-1 female germline stem cell (with Top235-1) Top217-6 imaginal disc | third instar larval stage | heat sensitive Top217-2 mesothoracic tergum, with Scer\GAL4pnr-MD237 Top2GD4570 mushroom body | somatic clone Top2unspecified ovary (with Top217-6) Top235-1 ovary (with Top235-1) Top217-6 polytene chromosome (with Df(2L)Exel9043) Top217-5 Top235-14
Classical Alleles ( 22 )
For All Classical Alleles Show
Allele of Top2 Class Mutagen Stocks Known lesion Top2LA00892 P-element activity 2 -- Top2c05388 piggyBac transposase 1 -- Top2d05357 P-element activity 1 -- Top2f05145 piggyBac transposase 1 -- Top2MB04073 1 -- Top217-7 amorphic allele - molecular evidence ethyl methanesulfonate 0 Yes Top235-14 amorphic allele - molecular evidence ethyl methanesulfonate 0 Yes Top217-1 ethyl methanesulfonate 0 Yes Top217-2 ethyl methanesulfonate 0 Yes Top217-3 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes Top217-5 ethyl methanesulfonate 0 Yes Top217-6 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes Top235-12 hypomorphic allele - genetic evidence, hypomorphic allele - molecular evidence ethyl methanesulfonate 0 Yes Top235-13 hypomorphic allele - genetic evidence, hypomorphic allele - molecular evidence ethyl methanesulfonate 0 Yes Top235-1 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes Top235-2 ethyl methanesulfonate 0 Yes Top235-3 hypomorphic allele - molecular evidence ethyl methanesulfonate 0 Yes Top235-4 ethyl methanesulfonate 0 Yes Top235-5 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes Top235-6 hypomorphic allele - genetic evidence, hypomorphic allele - molecular evidence ethyl methanesulfonate 0 Yes Top2Df17 P-element activity 0 Yes Top2unspecified 0 --
Alleles Carried on Transgenic Constructs ( 6 )
For All Alleles Carried on Transgenic Constructs Show
Allele of Top2 Class Mutagen Stocks Known lesion Top2GD4570 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes Top2GL00338 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes Top2JF01300 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes Top2+t7.1 in vitro construct - genomic fragment 0 Yes Top2dsRNA.cHa in vitro construct - RNAi 0 Yes Top2NIG.10223R in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2L)Exel7076 (Ryder, 2004.4.26) Df(2L)Exel8041 (Ryder, 2004.4.26) Df(2L)Top2Df35 (Hohl et al., 2012) Df(2L)TW158 (Palopoli and Wu, 1996)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD4570} NA P{NIG.10223R} NA P{TRiP.GL00338} NA P{TRiP.JF01300} NA characterization construct P{Top2.+7.1} NA
Insertions
	Type of insertions Name Expression data miscellaneous insertions P{}Top2Df17 NA PBac{PB}Top2c05388 NA PBac{WH}Top2f05145 NA insertion of mobile activating element P{Mae-UAS.6.11}Top2LA00892 NA P{XP}Top2d05357 NA insertion of enhancer trap binary system Mi{ET1}Top2MB04073 No
Gene Ontology: Function, Process & Cellular Component ( 25 unique terms )
Terms Based on Experimental Evidence ( 18 terms )
Molecular Function
CV term Evidence References
DNA binding inferred from direct assay (Rzepecki and Fisher, 2000) DNA topoisomerase type II (ATP-hydrolyzing) activity inferred from direct assay (Rzepecki and Fisher, 2000, Lee et al., 1989) four-way junction DNA binding inferred from direct assay (Strick et al., 2000, Zechiedrich and Osheroff, 1990) mRNA binding inferred from direct assay (Rzepecki and Fisher, 2000) rDNA binding inferred from direct assay (Blattes et al., 2006) satellite DNA binding inferred from direct assay (Blattes et al., 2006)
Biological Process
CV term Evidence References
chromatin silencing inferred from direct assay (Blattes et al., 2006) DNA topological change inferred from direct assay (Strick et al., 2000, Lee et al., 1989) metaphase plate congression inferred from direct assay (Chang et al., 2003) mitosis inferred from mutant phenotype (Ducat et al., 2008) mitotic chromosome condensation inferred from mutant phenotype (Goshima et al., 2007) mitotic G2 DNA damage checkpoint inferred from genetic interaction with nbs (Kondo and Perrimon, 2011) mitotic spindle organization inferred from mutant phenotype (Goshima et al., 2007) sister chromatid segregation inferred from direct assay (Chang et al., 2003, Coelho et al., 2003)
Cellular Component
CV term Evidence References
NOT ACF complex inferred from direct assay (Ito et al., 1997) CHRAC inferred from direct assay (Varga-Weisz et al., 1997) NOT CHRAC inferred from direct assay (Eberharter et al., 2001) chromosome inferred from direct assay (Zhang et al., 2000) nucleus inferred from direct assay (Colflesh et al., 1999, Zhang et al., 2000, Lee et al., 1989, Gemkow et al., 2001)
Terms Based on Predictions or Assertions ( 10 terms )
Molecular Function
CV term Evidence References
ATP binding inferred from electronic annotation with InterPro:IPR001154, InterPro:IPR001241, InterPro:IPR002205, InterPro:IPR003594, InterPro:IPR013506, InterPro:IPR013757, InterPro:IPR013758, InterPro:IPR013759, InterPro:IPR013760, InterPro:IPR018522 (FlyBase Curators et al., 2004-) DNA topoisomerase type II (ATP-hydrolyzing) activity non-traceable author statement (Swiss-Prot Project Members, 1990.4.1)
Biological Process
CV term Evidence References
DNA-dependent DNA replication inferred from biological aspect of ancestor with PANTHER:PTN000018846 (assigned by RefGenome) (Gaudet et al., 2010) meiosis traceable author statement (Rzepecki and Fisher, 2000) mitosis traceable author statement (Rzepecki and Fisher, 2000) mitotic recombination inferred from biological aspect of ancestor with PANTHER:PTN000018888 (assigned by RefGenome) (Gaudet et al., 2010) resolution of meiotic recombination intermediates inferred from biological aspect of ancestor with PANTHER:PTN000018888 (assigned by RefGenome) (Gaudet et al., 2010)
Cellular Component
CV term Evidence References
NOT nucleoid inferred from biological aspect of ancestor with PANTHER:PTN000018888 (assigned by RefGenome) (Gaudet et al., 2010) nucleus non-traceable author statement (Swiss-Prot Project Members, 1990.4.1) synaptonemal complex inferred from biological aspect of ancestor with PANTHER:PTN000018888 (assigned by RefGenome) (Gaudet et al., 2010)
Sequence Ontology: Class of Gene
	nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
protein-protein
Interacting group Assay References
Top2 - mod(mdg4) anti bait coimmunoprecipitation, western blot, two hybrid (Ramos et al., 2011) Top2 - scf anti bait coimmunoprecipitation, western blot (Kobayashi et al., 1998)
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions Top2 allele Gene References
External Data
Linkouts	DroID - A comprehensive database of gene and protein interactions. • FBgn0003732 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 35225
Orthologs
OrthoDB Orthologs (55) - 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 EOG670V2N Dipteran inclusive ortholog search EOG6FXRGH Insect inclusive ortholog search EOG6P2NX4 Arthropod inclusive ortholog search EOG6RR4Z6 Metazoa inclusive ortholog search EOG6RC3B9
Orthologs in Drosophila Species (EOG670V2N)
Organism Common Name Gene AAA Syntenic Ortholog Multiple Dmel Genes in this Orthologous Group Drosophila melanogaster fruit fly  Dmel\Top2     Drosophila simulans   Dsim\Top2 Y   Drosophila sechellia   Dsec\Top2 Y   Drosophila erecta   Dere\GG21630 Y   Drosophila yakuba   Dyak\GE12650 Y   Drosophila ananassae   Dana\GF15632 Y   Drosophila pseudoobscura pseudoobscura   Dpse\GA10169 Y   Drosophila persimilis   Dper\GL21103 Y   Drosophila willistoni   Dwil\GK24224 Y   Drosophila virilis   Dvir\GJ18099 Y   Drosophila mojavensis   Dmoj\GI19824 Y   Drosophila grimshawi   Dgri\GH10628 Y
Orthologs in non-Drosophila Dipterans (EOG6FXRGH)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Aedes aegypti Yellow fever mosquito  Aaeg\AAEL012584   Anopheles gambiae Malaria mosquito  Agam\AGAP008917   Culex quinquefasciatus Southern house mosquito  Cqui\CPIJ010018
Orthologs in non-Dipteran Insects (EOG6P2NX4)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Apis mellifera Western honey bee  Amel\GB50635   Nasonia vitripennis Parasitic wasp  Nvit\Nasvi2EG001287   Acromyrmex echinatior Panamanian leafcutter ant  Aech\AECH13560   Atta cephalotes Leafcutter ant  Acep\ACEP11747   Camponotus floridanus Florida carpenter ant  Cflo\CFLO13784   Harpegnathos saltator Jerdons jumping ant  Hsal\HSAL23213   Linepithema humile Argentine ant  Lhum\LH22942   Pogonomyrmex barbatus Red harvester ant  Pbar\PB26068   Solenopsis invicta Red fire ant  Sinv\SINV13620   Acyrthosiphon pisum Pea aphid  Apim\ACYPI007897   Acyrthosiphon pisum Pea aphid  Apim\ACYPI065069   Bombyx mori Silkmoth  Bmor\BGIBMGA011615   Pediculus humanus Human body louse  Phum\PHUM413330   Tribolium castaneum Red flour beetle  Tcas\Top2
Orthologs in non-Insect Arthropods (EOG6RR4Z6)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Daphnia pulex Water flea  Dpux\JGI_V11_329527   Daphnia pulex Water flea  Dpux\JGI_V11_274376   Daphnia pulex Water flea  Dpux\JGI_V11_47063   Daphnia pulex Water flea  Dpux\JGI_V11_333898   Daphnia pulex Water flea  Dpux\JGI_V11_329529   Daphnia pulex Water flea  Dpux\JGI_V11_323396   Daphnia pulex Water flea  Dpux\JGI_V11_332921   Ixodes scapularis Deer tick  Isca\ISCW016225
Orthologs in non-Arthropod Metazoa (EOG6RC3B9)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Caenorhabditis elegans Nematode  Cele\R05D3.12   Caenorhabditis elegans Nematode  Cele\K12D12.1   Caenorhabditis elegans Nematode  Cele\Y46H3C.4   Caenorhabditis elegans Nematode  Cele\cin-4   Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_018730gn   Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_027880gn   Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_010345gn   Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_004587gn   Danio rerio Zebrafish  Drer\top2b   Xenopus tropicalis Western clawed frog  Xtro\top2b   Xenopus tropicalis Western clawed frog  Xtro\top2a   Gallus gallus Domestic chicken  Ggal\TOP2B   Gallus gallus Domestic chicken  Ggal\TOP2A   Mus musculus House mouse  Mmus\ENSMUSG00000020914   Mus musculus House mouse  Mmus\Top2b   Rattus norvegicus Norway rat  Rnor\ENSRNOG00000032404   Homo sapiens Human  Hsap\ENSG00000077097   Homo sapiens Human  Hsap\TOP2A
Human Orthologs (2)
Gene OMIM HGNC Hsap\ENSG00000077097     Hsap\TOP2A
AAA Orthologs (11) based on analysis using Dmel annotation version 4.3
Organism Gene Drosophila simulans Dsim\Top2 Drosophila sechellia Dsec\Top2 Drosophila erecta Dere\GG21630 Drosophila yakuba Dyak\GE12650 Drosophila ananassae Dana\GF15632 Drosophila pseudoobscura pseudoobscura Dpse\GA10169 Drosophila persimilis Dper\GL21103 Drosophila willistoni Dwil\GK24224 Drosophila virilis Dvir\GJ18099 Drosophila mojavensis Dmoj\GI19824 Drosophila grimshawi Dgri\GH10628
Stocks & Reagents
Stocks Listed in FlyBase ( 9 )
Bloomington	24670 w1118; Mi{ET1}Top2MB04073 31342 y1 v1; P{TRiP.JF01300}attP2 35416 y1 sc* v1; P{TRiP.GL00338}attP2 22256 y1 w*; P{Mae-UAS.6.11}Top2LA00892
Harvard	- P{XP}Top2d05357 - PBac{PB}Top2c05388 - PBac{WH}Top2f05145
Kyoto	120892 y* w*; P{Mae-UAS.6.11}Top2LA00892
VDRC	v30625 w1118; P{GD4570}v30625
Genomic Clones ( 2 )
	BACR27M12 BACR28G24 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 135 )
	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	RE49802
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	AT29794 GH01096 GH09845 GH09857 GH11413 GM01217 GM04316 GM05438 GM07016 GM12611 GM16411 GM21208 GM22085 GM30444 LD01260 LD03057 LD04649 LD05895 LD06009 LD08127 LD08679 LD08941 LD09169 LD11135 LD11225 LD11893 LD12884 LD12928 LD14948 LD15786 LD17061 LD18896 LD19109 LD19521 LD19778 LD20660 LD21590 LD22377 LD23370 LD23503 LD26033 LD26479 LD29213 LD32256 LD34789 LD35208 LD35887 LD38976 LD39092 LD44963 LD46670 LD48039 LP13970 RE06939 RE22761 RE27317 RE29211 RE30402 RE32820 RE34062 RE49815 RE60669
Other clones	12174 21302 29619 30967 54518 63363 65452 66955 81712 104848 105194 153360 571156 CK01.11B.F8 CK01.41A.G11 CK01.41D.G11 CK01.61B.B12 CK01.70D.E10 Dmel_dig_018779_0715_1720 EC053114 EC053608 EC054092 EC056487 EK049140 EK055206 EK056251 EK063840 EK112836 EK139154 EK170602 EK173515 EK179748 EK191956 EK195117 EK229635 EK237002 EK257044 FGC023D11 FGM228E03 FGM312H12 LD20494 LD24716 LD28613 RM05081 RM05181 RM05381 RM05881 RM05981 RM06081 RM06281 RM06381 RM06481 RM06581 RP01002 RP01532 RP01591 RP001015999 RP001016544 RP001021975 RP001026051 RP001029098 RP001033445 RP001033802 RP001035235 RP001040158 RP001043559 RP001046658 RP001051401 RP001052106 RP001053925 RP001054676 UUGC0818
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0003732 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 35225
Antibody Information
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of
Additional comments
Other Comments
	RNAi screen using dsRNA made from templates generated with primers directed against this gene results in defects in chromosome condensation and chromosome misalignment on the metaphase spindle when assayed in S2 cells. This phenotype can be observed when the screen is performed with or without Cdc27 dsRNA. (Goshima et al., 2007) Expression is enriched in embryonic gonads. (Shigenobu et al., 2006) Area matching Drosophila Type II topoisomerase gene, Acc. No. X61209. (Davies, 2001.3.15) Amino acid residue Lys 359 is critical for the catalytic activity of Top2. (Hu et al., 1998) In a sample of 79 genes with multiple introns, 33 showed significant heterogeneity in G+C content among introns of the same gene and significant positive correspondence between the intron and the third codon position G+C content within genes. These results are consistent with selection adding against preferred codons at the start of genes. (Kliman and Eyre-Walker, 1998) The functional interaction domain for quinolones is mapped and the site of action is shown to overlap with the domain for etoposide, amsacrine and genistein. Results support the hypothesis that many DNA cleavage-enhancing drugs share a common interaction domain on the enzyme. (Elsea et al., 1997) Apurinic sites located within the base of the 5' overhang generated by Top2-mediated DNA scission cause levels of enzyme-associated DNA breaks to increase. Conversely apurinic sites located immediately outside the cleavage overhang inhibits DNA scission. Results indicate apurinic sites have a profound influence on the activity of Top2 enzyme and act as position-specific Top2 positions. (Kingma and Osheroff, 1997) Several forms of spontaneous DNA damage are capable of acting as endogenous poisons of Top2. (Kingma and Osheroff, 1997) A chromatin-accessibility complex (CHRAC), which uses energy to increase the general accessibility of DNA in chromatin, has been identified. CHRAC can also function during chromatin assembly, using ATP to convert irregular chromatin into a regular array of nucleosomes with even spacing. CHRAC contains 5 subunits, two of which have been identified as the Iswi and Top2 gene products. (Varga-Weisz et al., 1997) barr protein associates with Top2 protein and affects its activity, this suggests barr protein functions in chromatid decatenation during the metaphase-anaphase transition. (Bhat et al., 1996) Etoposide.Top2 interactions (rather than etoposide.DNA interactions) mediate enzyme-DNA cleavage complex formation. Maximal levels of etoposide-induced scission reflect the ability of the drug to inhibit religation at specific sequences rather than affinity of the drug to site-specific enzyme.DNA complexes. (Burden et al., 1996) Staurosporine inhibits catalytic activity of Top2 by blocking the transfer of phosphodiester bonds from DNA to the active tyrosine site. Other inhibitors most likely inactivate Top2 by alkylation of essential amino acids. (Lassota et al., 1996) Abasic sites are potent enhancers of double strand DNA cleavage mediated by Top2. Results suggest abasic sites represent cellular Top2 poisons and implies anticancer drugs mimic actions of these endogenous DNA lesions. (Kingma et al., 1995) Top2 is involved in protein-protein interactions that maintain nuclear localisation of Top2. Top2 has both RNA- and DNA-dependent anchorages in embryo nuclei. (Meller and Fisher, 1995) Topoisomerase 2 is distributed throughout interphase nuclei, except for the nucleoli, as judged by antibody staining. In some areas of nuclei localisation is not coincident with chromatin. (Meller et al., 1995) The ability of Top2 to cleave an oligonucleotide containing a hairpin has been studied. (Froelich-Ammon et al., 1994) Genetic and biochemical properties of a series of insertion linker mutations is used to analyse the domain structure of Top2. (Lee and Hsieh, 1994) Most Top2 in cell free extracts of 0-2hr embryos appears to be nonnuclear and soluble. This pool of Top2 is detectable in situ and can be detected in extracts from older (16-19hr) embryos and Kc tissue cells. These nonnuclear Top2 containing particles are composed largely of Top2 and an unknown RNA molecule(s). (Meller et al., 1994) Top2 is a Z-DNA binding protein, DNA binding characteristics are determined by filter binding and gel retardation. The Z-DNA binding activity of undegraded Top2 may be important in targetting the enzyme both to structural motifs required for chromatin organisation and to sites of local supercoiling. (Arndt-Jovin et al., 1993) Anti-Top2 antibodies or epipodophylotoxin VM26 injected into blastoderm embryos inhibits Top2 activity and prevents or hinders anaphase segregation of chromatids. High concentrations of inhibitors block chromosome condensation and the arrangement of chromosomes on the metaphase plate. (Buchenau et al., 1993) The effect of Pkc53E mediated phosphorylation on each step of Top2 catalytic cycle is analysed. As with Casein kinase II (CkIIβ and CkIIβ), stimulation of enzyme activity by Pkc53E correlates with an increased rate of Top2 catalysed ATP hydrolysis. Modification did not affect any step of the enzymes catalytic cycle that preceded the ATPase step. (Corbett et al., 1993) The effects of several drugs on the DNA strand passage event mediated by Top2 have been analysed. (Corbett et al., 1993) Top2-mediated DNA cleavage in the 5' region of an Hsp70 gene has been analysed. (Kroeger et al., 1993) Top2 is involved in different functions of the 1731 LTR. (Nahon et al., 1993) In defining the relationships between drug classes and their interaction domains on Top2 results indicate that there are mechanistic differences between etoposide and a number of other Top2-targeted antineoplastic agents and that the interaction domain of novobocin on the enzyme does not overlap those of several DNA cleavage-enhancing drugs. (Robinson et al., 1993) The distribution of microinjected fluorescently labelled Top2 has been monitored using time-lapse, three dimensional movies. Experiments show Top2 is localised to the nucleus and on chromosomes is spatially and temporally regulated. (Swedlow et al., 1993) The effects of casein kinase II-mediated phosphorylation on the individual steps of the enzymes catalytic cycle are characterised. Results strongly suggest that modification stimulates the activity of Top2 by enhancing the ability of the enzyme to hydrolyse its high energy ATP cofactor. (Corbett et al., 1992) To investigate the potential role for the passage helix in enzyme-mediated DNA cleavage, interactions between Top2 and a 40bp oligonucleotide that contains a specific enzyme recognition/cleavage sequence are characterised. Results indicate that the enzyme has to interact with more than a single double-stranded 40-mer in order for nucleic acid breakage to take place. (Corbett et al., 1992) Top2 can mediate illegitimate recombination events, at least in vitro, by a mechanism that does not rely on double-stranded DNA cleavage or subunit exchange. (Gale and Osheroff, 1992) The interaction of Top1 and Top2 gene products with transcriptionally active and inactive Act5C, Act57B and Hsp70B has been compared. Topoisomerase II binding to actin and Hsp70B sequences occurs on both transcriptionally active and inactive chromatin. An unusual type of topoisomerase II binding site was identified which is associated with the 5' ends of inactive Hsp70 and actin genes, suggesting that this enzyme may be associated with repression of gene transcription. (Kroeger and Rowe, 1992) Formation of DNA/Top2 complexes have been demonstrated using CsCl equilibrium density centrifugation: three types of complex that are in equilibrium with each other exist. The complex formed in the presence of the VM26 tumour suppressor is extremely stable. (Lee and Hsieh, 1992) 8-methoxycaffeine has been demonstrated to inhibit the activity of Top2 by interfering with the interaction of Top2 with DNA. (Tornaletti et al., 1992) Effects of short-wave UV-induced photoproducts on the enzymatic activity of Top2 are investigated. DNA relaxation reaction is inhibited and this correlates with an inhibition of the enzyme DNA strand passage event. Results suggest repair of cyclobutane pyrimidine dimers is important for the catalytic function of Top2. (Corbett et al., 1991) Top2 enzyme activity has been studied. (Glikin et al., 1991) EM is used to further characterise the Top2-DNA complex and to directly visualize the position of Top2 molecules on DNA containing sequence-directed bends. (Howard et al., 1991) The Top2 gene lies immediately adjacent to the location of the tandem duplication shown to be specific to RanGapSD chromosomes, and falls within the region from which several RanGapSD-related transcripts are encoded. However no obvious difference in Top2 transcription could be detected in RanGapSD mutant flies. (Powers and Ganetzky, 1991) Pre- and post-strand passage DNA cleavage complexes of Top2 serve as physiological targets for structurally disparate antineoplastic drugs. (Robinson and Osheroff, 1991) The effects of CP-67,804 and CP-115,953, two novel quinolone derivatives, on the enzymatic activity of Top2 are characterised. Both drugs enhance the pre- and poststrand passage DNA strand cleavage, neither inhibited the ability to religate cleaved DNA. Results strongly suggest CP-67,804 and CP-115,953 represent a novel class of Top2-targeted drugs. (Robinson et al., 1991) Top2 mediated cleavage of single-stranded DNA takes pace prior to the addition of SDS in an in vitro system. (Gale and Osheroff, 1990) Structurally disparate class of Top2 targeted antineoplastic drugs stabilise the enzyme's DNA cleavage complex primarily by interfering with the ability of Top2 to religate DNA. (Robinson and Osheroff, 1990) Top2 enzyme recognises supercoiled DNA at least in part by interacting preferentially with crossover points on DNA. (Zechiedrich and Osheroff, 1990) A number of anon-topo sites have been mapped and sequenced to study the sequence specificity of double strand DNA cleavage by Top2. (Sander and Hsieh, 1985) DNA molecules with topological knots are detected in a reaction mixture containing circular DNA and purified Top2 enzyme. This knotting reaction has been characterised. (Hsieh, 1983) Top2 protein has been purified and characterised. (Hsieh, 1983) The relaxation of negatively supercoiled circular DNA by purified Top2 enzyme has been studied. (Osheroff et al., 1983) The cleavage of double strand DNA by Top2 enzyme has been studied. (Sander and Hsieh, 1983) Top2 is an essential gene that encodes the large subunit of type II DNA topoisomerase, an enzyme believed to play an important role in the condensation, decondensation, and segregation of chromosomes. The enzyme is a major component of the nuclear matrix of Drosophila cells (Berrios, Osheroff and Fisher, 1985) and is distributed along polytene chromosomes paralleling the distribution of the DNA (Heller, Shelton, Dietrich, Elgin and Brutlag, 1986). It is believed to act by passing a DNA segment through a transient double-stranded break in another segment. Major cleavage sites for type II topoisomerase have been found in nontranscribed spacer segments and in the 5' and 3' ends of Hsp70 and the histone genes (Udvardy, Schedl, Sander and Hsieh, 1985). When prepared from embryos, the purified enzyme is made up of a major polypeptide encoded by Top2 of 166,000 daltons, with binding sites for both DNA and ATP, and, in addition, smaller polypeptides of 30,000-40,000 and 132,000-145,000 daltons (Sander and Hsieh, 1983; Shelton, Osheroff and Brutlag, 1983; Heller, Shelton, Dietrich, Elgin and Brutlag, 1986). Protein kinase activity is associated with Drosophila topoisomerase II (Sander, Nolan and Hsieh, 1984; Ackerman, Glover and Osheroff, 1985).
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_057412 NM_001273651 RefSeq (Proteins) • NP_476760 NP_001260580 DDBJ / EMBL / GenBank • AAF53802 AAQ23558 AC005428 AC007082 AGB93115 AX093881 BI364637 BT010240 CAA43523 CZ469933 CZ474486 CZ487766 DU708674 U53787 U53788 U53789 U53790 U53791 U53792 U53793 U53794 U53795 U53796 U53797 U53798 X61209 Entrez Gene - A searchable database of RefSeq genes. • 35225 UniProtKB/Swiss-Prot • P15348 UniProtKB/TrEMBL • Q6NR40
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • DNA topoisomerase, type IIA, conserved site (IPR018522) Toprim domain (IPR006171) DNA topoisomerase II, eukaryotic-type (IPR001154) DNA topoisomerase, type IIA (IPR001241) DNA topoisomerase, type IIA, subunit A/C-terminal (IPR002205) Histidine kinase-like ATPase, ATP-binding domain (IPR003594) DNA topoisomerase, type IIA, subunit B, domain 2 (IPR013506) DNA topoisomerase, type IIA, subunit A/ C-terminal, alpha-beta (IPR013758) DNA topoisomerase, type IIA, central domain (IPR013759) DNA topoisomerase, type IIA-like domain (IPR013760) Type IIA DNA topoisomerase subunit A, alpha-helical domain (IPR013757) Ribosomal protein S5 domain 2-type fold (IPR020568) Arginine repressor C-terminal-like domain (IPR024946) Ribosomal protein S5 domain 2-type fold, subgroup (IPR014721)
Linkouts
	DroID - A comprehensive database of gene and protein interactions. • FBgn0003732 DRSC - Results from RNAi screens. • FBgn0003732 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003732 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG10223-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0003732 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 35225 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 35225 modMine - Data generated by the modENCODE project. • FBgn0003732 REDfly - A database of transcriptional regulatory elements. • FBgn0003732
Synonyms & Secondary IDs ( 21 )
Reported As
Symbol Synonym	CG10223 (Whitfield, 2000.11.20, Whitfield, 2000.10.6, Przewloka et al., 2007, Goshima et al., 2007, Cakouros et al., 2008, Ducat et al., 2008, Ni et al., 2008, Ni et al., 2011.7.19, Cammarato et al., 2011) CHRAC (Fry and Peterson, 2001, Ridgway and Almouzni, 2000, Farkas et al., 2000, Varga-Weisz et al., 1997) dCHRAC (Boyer et al., 2000, Peterson, 2000) dTopoII (Saurin et al., 2001) top2 (Gemkow et al., 2001, Binaschi et al., 2000, Hu et al., 1998, Binaschi et al., 2000) TOP2 (Nitiss, 1998, Kondo and Perrimon, 2011) Top2 (Mukai et al., 2006, Shigenobu et al., 2006, Goshima et al., 2007, Williams et al., 2007, Robida et al., 2007, Ramos et al., 2011, Somma et al., 2008, Kallappagoudar et al., 2010, Schuldiner et al., 2008, Presgraves et al., 2009, Wang et al., 2013, Hohl et al., 2012, Schrider et al., 2011, Japanese National Institute of Genetics, 2012.5.21, Ferree and Prasad, 2012) TopII (Lyttle, 1993, Powers and Ganetzky, 1991) TopoII (Charvin et al., 2003, Langst and Becker, 2001, Kingston and Narlikar, 1999, Bjorklund et al., 1999, Csink and Henikoff, 1998, Philp, 1998, Plank et al., 2006, Ramos et al., 2011, Pimenta-Marques et al., 2008, Ferree and Barbash, 2009) topoII (Berrios et al., 2000, de Saint Phalle et al., 2002, Peterson, 2000, Kwok et al., 1999, Kirsch-Volders et al., 1998, Chang et al., 1998, Varga-Weisz et al., 1997, Foe et al., 1993, Swedlow et al., 1993, Whalen et al., 1991) topo II (Hu et al., 2002, Eberharter et al., 2001, Fisher et al., 2001, Strick et al., 2000, Rzepecki and Fisher, 2000, Wilson et al., 2000, Ito et al., 1999, Sikorav et al., 1998, Isaacs et al., 1998, Kobayashi et al., 1998, Bhat et al., 1996, Jongens et al., 1988, Hossain et al., 2004) TOPOII (Lupo et al., 2001) Topo II (Lupo et al., 1999, Przewloka et al., 2007, Chelouah et al., 2011, Orr et al., 2007, Chang et al., 2003, Wu et al., 2008, Oliveira et al., 2010, Pek and Kai, 2011) TOPO II (Coelho et al., 2008)
Name Synonym	DNA topoisomerase II (Binaschi et al., 2000, Savvidou et al., 2005) Top 2 (Schulte et al., 2011) Topoisomerase 2 (Mukai et al., 2006, Ducat et al., 2008, Nie et al., 2009, Cammarato et al., 2011) topoisomerase II (Rodriguez-Arnaiz et al., 2005, Geyer and Clark, 2002, Fortune et al., 2002, Gasser, 2002, Hirano, 2002, Kobayashi and Hirose, 1999, Udvardy, 1999, Kwok et al., 1998, Kwok and Hurley, 1998, Capranico and Binaschi, 1998, Burden and Osheroff, 1998, Sheridan et al., 1998, Fortune and Osheroff, 1998, Anderson et al., 1998, Alexiadis et al., 1998, Misteli and Spector, 1998, Jo and Topal, 1998, Austin and Marsh, 1998, Zink and Cremer, 1998, Gregory and Horz, 1998, Ito et al., 1997, Marx et al., 1997, Cline et al., 1997, Varga-Weisz et al., 1997, Ito et al., 1997, Marzano et al., 1997, Eder et al., 1993, Swedlow et al., 1992, Udvardy and Schedl, 1991, Swedlow et al., 1991, Somma et al., 2003, Elsea et al., 1997, Froelich-Ammon et al., 1994, Kroeger et al., 1993, Arndt-Jovin et al., 1993, Robinson et al., 1993, Corbett et al., 1993, Glikin et al., 1991, Robinson and Osheroff, 1991, Zechiedrich and Osheroff, 1990, Osheroff et al., 1983, Sander and Hsieh, 1983, Hsieh, 1983, Blattes et al., 2006, Mikhaylova et al., 2006, Gemkow et al., 2001, Lee et al., 1989, Williams et al., 2007, Binaschi et al., 2000, Cakouros et al., 2008, Chang et al., 2003, Coelho et al., 2003, Oliveira et al., 2010, Kondo and Perrimon, 2011) Topoisomerase II (Honore and Vorum, 2000, Shapiro et al., 1999, Sander and Hsieh, 1985, Somma et al., 2008, Orr et al., 2007, Pimenta-Marques et al., 2008, Ferree and Barbash, 2009) type II DNA topoisomerase (Stasiak, 2000) type II topoisomerase (Larson and Drummond, 2001, Hsieh, 1983)
Secondary FlyBase IDs
References ( 219 )
Generate a list of	All references relating to this gene ( 219 )
List References by type	Research paper  ( 148 ) Supplementary material  ( 5 ) Review  ( 31 ) Erratum  ( 1 ) Personal communication to FlyBase  ( 12 ) Abstract  ( 11 ) FlyBase analysis  ( 2 ) Conference report  ( 1 ) Stock list  ( 1 ) Patent  ( 1 ) DNA/RNA sequence record  ( 1 ) Protein sequence record  ( 1 ) Website  ( 1 ) Curated genome annotation  ( 2 ) Book  ( 1 )
Recent research papers ( 9 )
	Wang et al., 2013, Nat. Struct. Mol. Biol. 20(2): 202--209 Chromatin proteins captured by ChIP-mass spectrometry are linked to dosage compensation in Drosophila. [FBrf0220693] Crawford et al., 2012, Cell Death Differ. 19(12): 2040--2048 Conservation of caspase substrates across metazoans suggests hierarchical importance of signaling pathways over specific targets and cleavage site motifs in apoptosis. [FBrf0220233] Hohl et al., 2012, Genetics 192(3): 843--856 Restoration of topoisomerase 2 function by complementation of defective monomers in Drosophila. [FBrf0220101] Chelouah et al., 2011, PLoS ONE 6(8): e23597 An Integrated Drosophila Model System Reveals Unique Properties for F14512, a Novel Polyamine-Containing Anticancer Drug That Targets Topoisomerase II. [FBrf0214738] Kondo and Perrimon, 2011, Sci. Signal. 4(154): rs1 A Genome-Wide RNAi Screen Identifies Core Components of the G2-M DNA Damage Checkpoint. [FBrf0212700] Pek and Kai, 2011, Curr. Biol. 21(1): 39--44 A role for vasa in regulating mitotic chromosome condensation in Drosophila. [FBrf0212718] Ramos et al., 2011, PLoS ONE 6(1): e16562 DNA Topoisomerase II Modulates Insulator Function in Drosophila. [FBrf0213027] Schrider et al., 2011, Genome Res. 21(12): 2087--2095 Genome-wide analysis of retrogene polymorphisms in Drosophila melanogaster. [FBrf0216827] Schulte et al., 2011, PLoS ONE 6(8): e23841 High-Content Chemical and RNAi Screens for Suppressors of Neurotoxicity in a Huntington's Disease Model. [FBrf0215266] Show all research papers ...
Recent reviews ( 1 )
	Ferree and Prasad, 2012, Genet. Res. Int. 2012: 430136 How can satellite DNA divergence cause reproductive isolation? Let us count the chromosomal ways. [FBrf0218238] Show all reviews ...