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Gene Dmel\ben

General Information
Symbol	Dmel\ben	Species	D. melanogaster
Name	bendless	Annotation symbol	CG18319
Feature type	protein_coding_gene	FlyBase ID	FBgn0000173
Gene Model Status	Current	Stock availability	5 publicly available
Also Known As	ubc13, UbcD3
Genomic Location
Chromosome (arm)	X	Recombination map	1-45.0
Cytogenetic map	12D2-12D2	Sequence location	X:13,890,378..13,893,838 [+]
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 bendless is referred to in FlyBase by the symbol Dmel\ben (CG18319, FBgn0000173). It is a protein_coding_gene from Drosophila melanogaster. Based on sequence similarity, it is predicted to have molecular function: ubiquitin-protein ligase activity. There is experimental evidence that it is involved in the biological process: jump response; response to anesthetic; flight behavior; axonogenesis; axon target recognition; photoreceptor cell morphogenesis; grooming behavior. 25 alleles are reported. The phenotypes of these alleles are annotated with: organ system subdivision; external compound sense organ; peripheral nervous system; adult segment; nervous system; adult mesothoracic segment; multi-cell-component structure; indirect flight muscle; synaptic neuropil subdomain; rhabdomere R7. It has 5 annotated transcripts and 5 annotated polypeptides. Protein features are: Ubiquitin-conjugating enzyme, E2; Ubiquitin-conjugating enzyme, active site; Ubiquitin-conjugating enzyme/RWD-like. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of very high expression to a trough of moderately high expression. Peak expression observed within 00-06 hour embryonic stages, during early pupal stages. Summary of FlyAtlas Anatomical Expression Data: High or moderate levels of expression observed in all larval and adult organs/tissues. Expression at high levels in the following post-embryonic organs or tissues: adult head, adult eye, larval/adult central nervous system, adult crop, larval/adult midgut, larval/adult hindgut, larval/adult Malpighian tubules, adult heart, larval/adult fat body, larval salivary gland, larval trachea, adult spermathecae, adult male accessory gland, larval carcass. Expression at moderate levels in the following post-embryonic organs or tissues: adult salivary gland, adult ovary, adult testis, adult carcass. Comments on Affy2 ProbeSet: ProbeSet 1641279_at completely aligns to an exonic region of only a subset of 1 of the 2 FlyBase-annotated transcript isoforms of ben. Gene sequence location is X:13890378..13893838.
User Contributed Data
	Beta version, feedback welcome
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	ben: bendless (J. Hall; R. Wyman) Adults have an aberrant startle response; they do not jump when presented with a lights-off stimulus. The cervical giant fiber, a brain neuron, has abnormal morphology. The normal giant fiber terminates in the thorax at a synapse onto the motoneuron innervating the tergo-trochanteral (TT) muscle (= jump muscle). In ben, this synapse is abnormal or absent. The normal lateral bend of the giant fiber toward the TT motoneuron in the mesothoracic neuromere is absent; the axon usually terminates at the midline with fine branches extending from its tip. Latency of response of the TT muscle to stimulation of the giant fiber is abnormally long, and muscles cannot follow stimulation at rates above 5 Hz. Rhabdomeres of the photoreceptor cells in the ommatidia heart shaped in cross section with indentation centrally oriented rather than round as is normal. Furthermore, the axons from photoreceptor cells R7 and R8 fail to make the right-angle turn into the optic medulla after traversing the lamina (Benzer). ben flies choose visible over ultraviolet wave lengths whereas wild-type flies make the opposite choice (Benzer).
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
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 12D2-12D2   Limits computationally determined from genome sequence between P{EP}EP1595EP1595 and P{EP}l(1)G0007EP764
Experimentally Determined Cytological Location
Cytogenetic map Notes References 12D-12D   (determined by in situ hybridisation)   (Matuschewski et al., 1996)
Experimentally Determined Recombination Data
Location	1-45.8 (Oh et al., 1994, Tanouye et al., 1987) 1-45.0
Left of (cM)	na (Tanouye et al., 1986, Oh et al., 1994) f (Thomas and Wyman, 1984)
Right of (cM)	v (Thomas and Wyman, 1984) g (Tanouye et al., 1986, Tanouye et al., 1987, Oh et al., 1994) P{ry1}SteR301.2 (Muralidhar and Thomas, 1993)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\ben for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
	Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated Low-frequency RNA-Seq exon junction(s) not annotated. Annotated transcripts do not represent all supported alternative splices within 5' UTR. Gene model reviewed during 5.45 Gene model reviewed during 5.50
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) ben-RA FBtr0073855  NM_078595   1890   151 ben-RB FBtr0300566  NM_001169281   852   151 ben-RC FBtr0307296  NM_001258734   2003   151 ben-RD FBtr0332843  NM_001272611   1852   151 ben-RE FBtr0332844  NM_001272612   1810   151
Additional Transcript Data & Comments
Reported size (kB)	2.0, 1.2 (northern blot) (Muralidhar and Thomas, 1993)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein ben-PA   FBpp0073686   17.2   151   5.28     NP_511150   AAF48338 ben-PB   FBpp0289793   17.2   151   5.28     NP_001162752   ACZ95287 ben-PC   FBpp0298297   17.2   151   5.28     NP_001245663   AFH07377 ben-PD   FBpp0305066   17.2   151   5.28     NP_001259540   AGB95382 ben-PE   FBpp0305067   17.2   151   5.28     NP_001259541   AGB95383
Additional Polypeptide Data & Comments
Reported size (kDa)	151 (aa) (Muralidhar and Thomas, 1993)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Ubiquitin-conjugating enzyme, E2 (IPR000608) Ubiquitin-conjugating enzyme/RWD-like (IPR016135) Ubiquitin-conjugating enzyme, active site (IPR023313)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA439856   AA441407   AA695402   AA820626   AA951369   AA978689   AE014298 AAF48338   AE014298 ACZ95287   AE014298 AFH07377   AE014298 AGB95382   AE014298 AGB95383   AI063934   AI109621   AI113934   AI133913   AI403213   AI455478   AX777388   AY069527 AAL39672   AY826157   BI171424   BI237077   BI237858   BI238647   BI239353   BI243079   BI358686   BI363449   BI366913   BI375734   BI482753   BI573246   BI573247   BI635003   BP540236   BP540238   BP546994   BP552010   BQ103174   CA806848   CA807109   CA807279   CO177062   CO185840   CO262532   CO266071   CO266638   CO266778   CO268256   CO269882   CO271313   CO271731   CO273464   CO277372   CO278944   CO279211   CO280375   CO281514   CO285451   CO287182   CO288045   CO290927   CO291359   CO291981   CO292516   CO293014   CO295456   CO303250   CO309591   CO312144   CO318167   CO330967   CO332838   CO332857   CO334080   CO337166   CO339828   CO340117   CO344429   EC051299   EC058224   EC059517   EC060905   EC061239   EC063686   EC064207   EC071339   EC071629   EC071977   EC073306   EC079015   EC080184   EC080327   EC084525   EC087737   EC088125   EC089612   EC090834   EC194927   EC196499   EC196520   EC201939   EC210147   EC214113   EC214136   EC214414   EC216652   EC218508   EC219001   EC221858   EC221983   EC224058   EC224121   EC228079   EC229076   EC230022   EC232264   EC233914   EC236740   EC239288   EC245987   EC251178   EC252028   EC254591   EC258706   EC264273   EL873520   EL873532   EL875571   EV586464   HQ383939   L20126 AAA28392   S70118 AAB30753
UniProtKB/Swiss-Prot	P35128
UniProtKB/TrEMBL	E1JJN1
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 | intense & ubiquitous   (Fisher et al., 2012) Comment:maternally deposited embryonic stage 4 -- 6   organism | intense & ubiquitous   (Fisher et al., 2012) embryonic stage 7 -- 8   organism | intense & ubiquitous   (Fisher et al., 2012) embryonic stage 9 -- 10   anterior endoderm   (Fisher et al., 2012) organism | ubiquitous   (Fisher et al., 2012) posterior endoderm   (Fisher et al., 2012) ventral nerve cord primordium   (Fisher et al., 2012) embryonic stage 11 -- 12   anterior midgut primordium   (Fisher et al., 2012) central brain primordium   (Fisher et al., 2012) organism | faint & ubiquitous   (Fisher et al., 2012) posterior midgut primordium   (Fisher et al., 2012) ventral nerve cord primordium   (Fisher et al., 2012) visceral muscle primordium   (Fisher et al., 2012) embryonic stage 11 -- 17   peripheral nervous system   (Muralidhar and Thomas, 1993) presumptive embryonic/larval central nervous system | presumptive   (Muralidhar and Thomas, 1993) embryonic stage 13 -- 16   embryonic brain   (Fisher et al., 2012) embryonic/larval midgut   (Fisher et al., 2012) embryonic/larval visceral muscle   (Fisher et al., 2012) organism | ubiquitous   (Fisher et al., 2012) ventral nerve cord   (Fisher et al., 2012) third instar larval stage   embryonic/larval muscle system   (Muralidhar and Thomas, 1993) embryonic/larval salivary gland   (Muralidhar and Thomas, 1993) subesophageal ganglion   (Muralidhar and Thomas, 1993) embryonic/larval digestive system   (Muralidhar and Thomas, 1993) imaginal disc   (Muralidhar and Thomas, 1993) ventral nerve cord   (Muralidhar and Thomas, 1993) proventriculus   (Muralidhar and Thomas, 1993) supraesophageal ganglion   (Muralidhar and Thomas, 1993) pupal stage & adult stage   supraesophageal ganglion   (Muralidhar and Thomas, 1993) thoracico-abdominal ganglion   (Muralidhar and Thomas, 1993) subesophageal ganglion   (Muralidhar and Thomas, 1993) retina   (Muralidhar and Thomas, 1993)
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
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: High or moderate levels of expression observed in all larval and adult organs/tissues. Expression at high levels in the following post-embryonic organs or tissues: adult head, adult eye, larval/adult central nervous system, adult crop, larval/adult midgut, larval/adult hindgut, larval/adult Malpighian tubules, adult heart, larval/adult fat body, larval salivary gland, larval trachea, adult spermathecae, adult male accessory gland, larval carcass. Expression at moderate levels in the following post-embryonic organs or tissues: adult salivary gland, adult ovary, adult testis, 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    1376.675 Larval Midgut    1217 Larval Hindgut    1409.8 Larval Malpighian Tubules    1612.4 Larval Fat Body    1033.7 Larval Salivary Gland    907.9 Larval Trachea    678.9 Larval Carcass    1082.475 Adult Head    1015 Adult Eye    1068.575 Adult Brain    996.3 Adult Thoracic-Abdominal Ganglion    1092.8 Adult Crop    1040.4 Adult Midgut    515.1 Adult Hindgut    801.3 Adult Malpighian Tubules    676.8 Adult Fat Body    984.5 Adult Salivary Gland    478.6 Adult Heart    518.65 Adult VirginFemale Spermatheca    784.5 Adult InseminatedFemale Spermatheca    842 Adult Ovary    221.1 Adult Testis    123 Adult Male Accessory Gland    629.5 Adult Carcass    481 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System  (1376.675) Larval Midgut  (1217) Larval Hindgut  (1409.8) Larval Malpighian Tubules  (1612.4) Larval Fat Body  (1033.7) Larval Salivary Gland  (907.9) Larval Trachea  (678.9) Larval Carcass  (1082.475) Adult Head  (1015) Adult Eye  (1068.575) Adult Brain  (996.3) Adult Thoracic-Abdominal Ganglion  (1092.8) Adult Crop  (1040.4) Adult Midgut  515.1 Adult Hindgut  (801.3) Adult Malpighian Tubules  (676.8) Adult Fat Body  (984.5) Adult Salivary Gland    478.6 Adult Heart  518.65 Adult VirginFemale Spermatheca  (784.5) Adult InseminatedFemale Spermatheca  (842) Adult Ovary    221.1 Adult Testis    123 Adult Male Accessory Gland  (629.5) Adult Carcass    481 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System  (1376.675) Larval Midgut  (1217) Larval Hindgut  (1409.8) Larval Malpighian Tubules  (1612.4) Larval Fat Body  1033.7 Larval Salivary Gland    907.9 Larval Trachea    678.9 Larval Carcass  (1082.475) Adult Head  1015 Adult Eye  1068.575 Adult Brain    996.3 Adult Thoracic-Abdominal Ganglion  (1092.8) Adult Crop  1040.4 Adult Midgut    515.1 Adult Hindgut    801.3 Adult Malpighian Tubules    676.8 Adult Fat Body    984.5 Adult Salivary Gland    478.6 Adult Heart    518.65 Adult VirginFemale Spermatheca    784.5 Adult InseminatedFemale Spermatheca    842 Adult Ovary    221.1 Adult Testis    123 Adult Male Accessory Gland    629.5 Adult Carcass    481 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    1376.675 Larval Midgut    1217 Larval Hindgut    1409.8 Larval Malpighian Tubules    1612.4 Larval Fat Body    1033.7 Larval Salivary Gland    907.9 Larval Trachea    678.9 Larval Carcass    1082.475 Adult Head    1015 Adult Eye    1068.575 Adult Brain    996.3 Adult Thoracic-Abdominal Ganglion    1092.8 Adult Crop    1040.4 Adult Midgut    515.1 Adult Hindgut    801.3 Adult Malpighian Tubules    676.8 Adult Fat Body    984.5 Adult Salivary Gland    478.6 Adult Heart    518.65 Adult VirginFemale Spermatheca    784.5 Adult InseminatedFemale Spermatheca    842 Adult Ovary    221.1 Adult Testis    123 Adult Male Accessory Gland    629.5 Adult Carcass    481 Expression Level Scale  Very high  log, scaled to maximum expression level Tissue   Expression Level Larval Central Nervous System    1376.675 Larval Midgut    1217 Larval Hindgut    1409.8 Larval Malpighian Tubules    1612.4 Larval Fat Body    1033.7 Larval Salivary Gland    907.9 Larval Trachea    678.9 Larval Carcass    1082.475 Adult Head    1015 Adult Eye    1068.575 Adult Brain    996.3 Adult Thoracic-Abdominal Ganglion    1092.8 Adult Crop    1040.4 Adult Midgut    515.1 Adult Hindgut    801.3 Adult Malpighian Tubules    676.8 Adult Fat Body    984.5 Adult Salivary Gland    478.6 Adult Heart    518.65 Adult VirginFemale Spermatheca    784.5 Adult InseminatedFemale Spermatheca    842 Adult Ovary    221.1 Adult Testis    123 Adult Male Accessory Gland    629.5 Adult Carcass    481 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System  (1376.675) Larval Midgut  (1217) Larval Hindgut  (1409.8) Larval Malpighian Tubules  (1612.4) Larval Fat Body  (1033.7) Larval Salivary Gland  (907.9) Larval Trachea  678.9 Larval Carcass  (1082.475) Adult Head  (1015) Adult Eye  (1068.575) Adult Brain  (996.3) Adult Thoracic-Abdominal Ganglion  (1092.8) Adult Crop  (1040.4) Adult Midgut  515.1 Adult Hindgut  801.3 Adult Malpighian Tubules  676.8 Adult Fat Body  (984.5) Adult Salivary Gland    478.6 Adult Heart  518.65 Adult VirginFemale Spermatheca  784.5 Adult InseminatedFemale Spermatheca  (842) Adult Ovary    221.1 Adult Testis    123 Adult Male Accessory Gland  629.5 Adult Carcass    481 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System  1376.675 Larval Midgut  1217 Larval Hindgut  1409.8 Larval Malpighian Tubules  1612.4 Larval Fat Body  1033.7 Larval Salivary Gland    907.9 Larval Trachea    678.9 Larval Carcass  1082.475 Adult Head  1015 Adult Eye  1068.575 Adult Brain    996.3 Adult Thoracic-Abdominal Ganglion  1092.8 Adult Crop  1040.4 Adult Midgut    515.1 Adult Hindgut    801.3 Adult Malpighian Tubules    676.8 Adult Fat Body    984.5 Adult Salivary Gland    478.6 Adult Heart    518.65 Adult VirginFemale Spermatheca    784.5 Adult InseminatedFemale Spermatheca    842 Adult Ovary    221.1 Adult Testis    123 Adult Male Accessory Gland    629.5 Adult Carcass    481 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    1376.675 Larval Midgut    1217 Larval Hindgut    1409.8 Larval Malpighian Tubules    1612.4 Larval Fat Body    1033.7 Larval Salivary Gland    907.9 Larval Trachea    678.9 Larval Carcass    1082.475 Adult Head    1015 Adult Eye    1068.575 Adult Brain    996.3 Adult Thoracic-Abdominal Ganglion    1092.8 Adult Crop    1040.4 Adult Midgut    515.1 Adult Hindgut    801.3 Adult Malpighian Tubules    676.8 Adult Fat Body    984.5 Adult Salivary Gland    478.6 Adult Heart    518.65 Adult VirginFemale Spermatheca    784.5 Adult InseminatedFemale Spermatheca    842 Adult Ovary    221.1 Adult Testis    123 Adult Male Accessory Gland    629.5 Adult Carcass    481 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    1015   NA  Eye    1068.575   NA  Brain    996.3   1376.675  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    1092.8   NA  Crop    1040.4   1217  Midgut    515.1   1409.8  Hindgut    801.3   1612.4  Malpighian Tubules    676.8   1033.7  Fat Body    984.5   907.9  Salivary Gland    478.6   NA  Heart    518.65   678.9  Trachea    NA   NA  VirginFemale Spermatheca    784.5   NA  InseminatedFemale Spermatheca    842   NA  Ovary    221.1   NA  Testis    123   NA  Male Accessory Gland    629.5   1082.475  Carcass    481 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    81 central nervous system, larvae L3    141 central nervous system, pupae P8    79 head, virgin 1-day female    60 head, virgin 4-day female    57 head, virgin 20-day female    53 head, mated 1-day female    49 head, mated 4-day female    58 head, mated 20-day female    39 head, mated 1-day male    68 head, mated 4-day male    88 head, mated 20-day male    82 salivary gland, larvae L3 wandering    74 salivary gland, white prepupae    209 digestive system, larvae L3 wandering    248 digestive system, 1-day adult    62 digestive system, 4-day adult    52 digestive system, 20-day adult    57 fat body, larvae L3 wandering    38 fat body, white prepupae    88 fat body, pupae P8    45 carcass, larvae L3 wandering    114 carcass, 1-day adult    35 carcass, 4-day adult    40 carcass, 20-day adult    34 ovary, virgin 4-day female    118 ovary, mated 4-day female    100 testis, mated 4-day male    19 accessory gland, mated 4-day male    17 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  (81) central nervous system, larvae L3  (141) central nervous system, pupae P8  (79) head, virgin 1-day female  (60) head, virgin 4-day female  (57) head, virgin 20-day female  (53) head, mated 1-day female  (49) head, mated 4-day female  (58) head, mated 20-day female  (39) head, mated 1-day male  (68) head, mated 4-day male  (88) head, mated 20-day male  (82) salivary gland, larvae L3 wandering  (74) salivary gland, white prepupae  (209) digestive system, larvae L3 wandering  (248) digestive system, 1-day adult  (62) digestive system, 4-day adult  (52) digestive system, 20-day adult  (57) fat body, larvae L3 wandering  (38) fat body, white prepupae  (88) fat body, pupae P8  (45) carcass, larvae L3 wandering  (114) carcass, 1-day adult  (35) carcass, 4-day adult  (40) carcass, 20-day adult  (34) ovary, virgin 4-day female  (118) ovary, mated 4-day female  (100) testis, mated 4-day male    19 accessory gland, mated 4-day male    17 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Tissue   Expression Level imaginal disc, larvae L3 wandering    81 central nervous system, larvae L3  (141) central nervous system, pupae P8    79 head, virgin 1-day female    60 head, virgin 4-day female    57 head, virgin 20-day female    53 head, mated 1-day female    49 head, mated 4-day female    58 head, mated 20-day female    39 head, mated 1-day male    68 head, mated 4-day male    88 head, mated 20-day male    82 salivary gland, larvae L3 wandering    74 salivary gland, white prepupae  (209) digestive system, larvae L3 wandering  (248) digestive system, 1-day adult    62 digestive system, 4-day adult    52 digestive system, 20-day adult    57 fat body, larvae L3 wandering    38 fat body, white prepupae    88 fat body, pupae P8    45 carcass, larvae L3 wandering  (114) carcass, 1-day adult    35 carcass, 4-day adult    40 carcass, 20-day adult    34 ovary, virgin 4-day female  (118) ovary, mated 4-day female    100 testis, mated 4-day male    19 accessory gland, mated 4-day male    17 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    81 central nervous system, larvae L3    141 central nervous system, pupae P8    79 head, virgin 1-day female    60 head, virgin 4-day female    57 head, virgin 20-day female    53 head, mated 1-day female    49 head, mated 4-day female    58 head, mated 20-day female    39 head, mated 1-day male    68 head, mated 4-day male    88 head, mated 20-day male    82 salivary gland, larvae L3 wandering    74 salivary gland, white prepupae    209 digestive system, larvae L3 wandering    248 digestive system, 1-day adult    62 digestive system, 4-day adult    52 digestive system, 20-day adult    57 fat body, larvae L3 wandering    38 fat body, white prepupae    88 fat body, pupae P8    45 carcass, larvae L3 wandering    114 carcass, 1-day adult    35 carcass, 4-day adult    40 carcass, 20-day adult    34 ovary, virgin 4-day female    118 ovary, mated 4-day female    100 testis, mated 4-day male    19 accessory gland, mated 4-day male    17 Expression Level Scale  Extremely high  log, scaled to maximum expression level Tissue   Expression Level imaginal disc, larvae L3 wandering    81 central nervous system, larvae L3    141 central nervous system, pupae P8    79 head, virgin 1-day female    60 head, virgin 4-day female    57 head, virgin 20-day female    53 head, mated 1-day female    49 head, mated 4-day female    58 head, mated 20-day female    39 head, mated 1-day male    68 head, mated 4-day male    88 head, mated 20-day male    82 salivary gland, larvae L3 wandering    74 salivary gland, white prepupae    209 digestive system, larvae L3 wandering    248 digestive system, 1-day adult    62 digestive system, 4-day adult    52 digestive system, 20-day adult    57 fat body, larvae L3 wandering    38 fat body, white prepupae    88 fat body, pupae P8    45 carcass, larvae L3 wandering    114 carcass, 1-day adult    35 carcass, 4-day adult    40 carcass, 20-day adult    34 ovary, virgin 4-day female    118 ovary, mated 4-day female    100 testis, mated 4-day male    19 accessory gland, mated 4-day male    17 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  (81) central nervous system, larvae L3  (141) central nervous system, pupae P8  (79) head, virgin 1-day female  (60) head, virgin 4-day female  (57) head, virgin 20-day female  (53) head, mated 1-day female  (49) head, mated 4-day female  (58) head, mated 20-day female  (39) head, mated 1-day male  (68) head, mated 4-day male  (88) head, mated 20-day male  (82) salivary gland, larvae L3 wandering  (74) salivary gland, white prepupae  (209) digestive system, larvae L3 wandering  (248) digestive system, 1-day adult  (62) digestive system, 4-day adult  (52) digestive system, 20-day adult  (57) fat body, larvae L3 wandering  (38) fat body, white prepupae  (88) fat body, pupae P8  (45) carcass, larvae L3 wandering  (114) carcass, 1-day adult  (35) carcass, 4-day adult  (40) carcass, 20-day adult  (34) ovary, virgin 4-day female  (118) ovary, mated 4-day female  (100) testis, mated 4-day male    19 accessory gland, mated 4-day male    17 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Tissue   Expression Level imaginal disc, larvae L3 wandering    81 central nervous system, larvae L3  141 central nervous system, pupae P8    79 head, virgin 1-day female    60 head, virgin 4-day female    57 head, virgin 20-day female    53 head, mated 1-day female    49 head, mated 4-day female    58 head, mated 20-day female    39 head, mated 1-day male    68 head, mated 4-day male    88 head, mated 20-day male    82 salivary gland, larvae L3 wandering    74 salivary gland, white prepupae  (209) digestive system, larvae L3 wandering  (248) digestive system, 1-day adult    62 digestive system, 4-day adult    52 digestive system, 20-day adult    57 fat body, larvae L3 wandering    38 fat body, white prepupae    88 fat body, pupae P8    45 carcass, larvae L3 wandering  114 carcass, 1-day adult    35 carcass, 4-day adult    40 carcass, 20-day adult    34 ovary, virgin 4-day female  118 ovary, mated 4-day female    100 testis, mated 4-day male    19 accessory gland, mated 4-day male    17 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    81 central nervous system, larvae L3    141 central nervous system, pupae P8    79 head, virgin 1-day female    60 head, virgin 4-day female    57 head, virgin 20-day female    53 head, mated 1-day female    49 head, mated 4-day female    58 head, mated 20-day female    39 head, mated 1-day male    68 head, mated 4-day male    88 head, mated 20-day male    82 salivary gland, larvae L3 wandering    74 salivary gland, white prepupae    209 digestive system, larvae L3 wandering    248 digestive system, 1-day adult    62 digestive system, 4-day adult    52 digestive system, 20-day adult    57 fat body, larvae L3 wandering    38 fat body, white prepupae    88 fat body, pupae P8    45 carcass, larvae L3 wandering    114 carcass, 1-day adult    35 carcass, 4-day adult    40 carcass, 20-day adult    34 ovary, virgin 4-day female    118 ovary, mated 4-day female    100 testis, mated 4-day male    19 accessory gland, mated 4-day male    17 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 moderately high expression. Peak expression observed within 00-06 hour embryonic stages, during early pupal 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    112 embryo 02-04hr    147 embryo 04-06hr    79 embryo 06-08hr    60 embryo 08-10hr    76 embryo 10-12hr    62 embryo 12-14hr    54 embryo 14-16hr    55 embryo 16-18hr    41 embryo 18-20hr    32 embryo 20-22hr    34 embryo 22-24hr    38 larva L1    34 larva L2    40 larva L3 12hr old    36 larva L3 puffstage 1-2    53 larva L3 puffstage 3-6    84 larva L3 puffstage 7-9    118 white prepupae new    142 white prepupae 12hr    97 white prepupae 24hr    77 pupae 2d postWPP    51 pupae 3d postWPP    36 pupae 4d postWPP    32 adult male 01day    35 adult male 05day    32 adult male 30day    29 adult female 01day    58 adult female 05day    79 adult female 30day    72 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (112) embryo 02-04hr  (147) embryo 04-06hr  (79) embryo 06-08hr  (60) embryo 08-10hr  (76) embryo 10-12hr  (62) embryo 12-14hr  (54) embryo 14-16hr  (55) embryo 16-18hr  (41) embryo 18-20hr  (32) embryo 20-22hr  (34) embryo 22-24hr  (38) larva L1  (34) larva L2  (40) larva L3 12hr old  (36) larva L3 puffstage 1-2  (53) larva L3 puffstage 3-6  (84) larva L3 puffstage 7-9  (118) white prepupae new  (142) white prepupae 12hr  (97) white prepupae 24hr  (77) pupae 2d postWPP  (51) pupae 3d postWPP  (36) pupae 4d postWPP  (32) adult male 01day  (35) adult male 05day  (32) adult male 30day  (29) adult female 01day  (58) adult female 05day  (79) adult female 30day  (72) Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr  (112) embryo 02-04hr  (147) embryo 04-06hr    79 embryo 06-08hr    60 embryo 08-10hr    76 embryo 10-12hr    62 embryo 12-14hr    54 embryo 14-16hr    55 embryo 16-18hr    41 embryo 18-20hr    32 embryo 20-22hr    34 embryo 22-24hr    38 larva L1    34 larva L2    40 larva L3 12hr old    36 larva L3 puffstage 1-2    53 larva L3 puffstage 3-6    84 larva L3 puffstage 7-9  (118) white prepupae new  (142) white prepupae 12hr    97 white prepupae 24hr    77 pupae 2d postWPP    51 pupae 3d postWPP    36 pupae 4d postWPP    32 adult male 01day    35 adult male 05day    32 adult male 30day    29 adult female 01day    58 adult female 05day    79 adult female 30day    72 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    112 embryo 02-04hr    147 embryo 04-06hr    79 embryo 06-08hr    60 embryo 08-10hr    76 embryo 10-12hr    62 embryo 12-14hr    54 embryo 14-16hr    55 embryo 16-18hr    41 embryo 18-20hr    32 embryo 20-22hr    34 embryo 22-24hr    38 larva L1    34 larva L2    40 larva L3 12hr old    36 larva L3 puffstage 1-2    53 larva L3 puffstage 3-6    84 larva L3 puffstage 7-9    118 white prepupae new    142 white prepupae 12hr    97 white prepupae 24hr    77 pupae 2d postWPP    51 pupae 3d postWPP    36 pupae 4d postWPP    32 adult male 01day    35 adult male 05day    32 adult male 30day    29 adult female 01day    58 adult female 05day    79 adult female 30day    72 Expression Level Scale  Extremely high  log, scaled to maximum expression level Developmental Stage   Expression Level embryo 00-02hr    112 embryo 02-04hr    147 embryo 04-06hr    79 embryo 06-08hr    60 embryo 08-10hr    76 embryo 10-12hr    62 embryo 12-14hr    54 embryo 14-16hr    55 embryo 16-18hr    41 embryo 18-20hr    32 embryo 20-22hr    34 embryo 22-24hr    38 larva L1    34 larva L2    40 larva L3 12hr old    36 larva L3 puffstage 1-2    53 larva L3 puffstage 3-6    84 larva L3 puffstage 7-9    118 white prepupae new    142 white prepupae 12hr    97 white prepupae 24hr    77 pupae 2d postWPP    51 pupae 3d postWPP    36 pupae 4d postWPP    32 adult male 01day    35 adult male 05day    32 adult male 30day    29 adult female 01day    58 adult female 05day    79 adult female 30day    72 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (112) embryo 02-04hr  (147) embryo 04-06hr  (79) embryo 06-08hr  (60) embryo 08-10hr  (76) embryo 10-12hr  (62) embryo 12-14hr  (54) embryo 14-16hr  (55) embryo 16-18hr  (41) embryo 18-20hr  32 embryo 20-22hr  (34) embryo 22-24hr  (38) larva L1  (34) larva L2  (40) larva L3 12hr old  (36) larva L3 puffstage 1-2  (53) larva L3 puffstage 3-6  (84) larva L3 puffstage 7-9  (118) white prepupae new  (142) white prepupae 12hr  (97) white prepupae 24hr  (77) pupae 2d postWPP  (51) pupae 3d postWPP  (36) pupae 4d postWPP  32 adult male 01day  (35) adult male 05day  32 adult male 30day  29 adult female 01day  (58) adult female 05day  (79) adult female 30day  (72) Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr  112 embryo 02-04hr  (147) embryo 04-06hr    79 embryo 06-08hr    60 embryo 08-10hr    76 embryo 10-12hr    62 embryo 12-14hr    54 embryo 14-16hr    55 embryo 16-18hr    41 embryo 18-20hr    32 embryo 20-22hr    34 embryo 22-24hr    38 larva L1    34 larva L2    40 larva L3 12hr old    36 larva L3 puffstage 1-2    53 larva L3 puffstage 3-6    84 larva L3 puffstage 7-9  118 white prepupae new  142 white prepupae 12hr    97 white prepupae 24hr    77 pupae 2d postWPP    51 pupae 3d postWPP    36 pupae 4d postWPP    32 adult male 01day    35 adult male 05day    32 adult male 30day    29 adult female 01day    58 adult female 05day    79 adult female 30day    72 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    112 embryo 02-04hr    147 embryo 04-06hr    79 embryo 06-08hr    60 embryo 08-10hr    76 embryo 10-12hr    62 embryo 12-14hr    54 embryo 14-16hr    55 embryo 16-18hr    41 embryo 18-20hr    32 embryo 20-22hr    34 embryo 22-24hr    38 larva L1    34 larva L2    40 larva L3 12hr old    36 larva L3 puffstage 1-2    53 larva L3 puffstage 3-6    84 larva L3 puffstage 7-9    118 white prepupae new    142 white prepupae 12hr    97 white prepupae 24hr    77 pupae 2d postWPP    51 pupae 3d postWPP    36 pupae 4d postWPP    32 adult male 01day    35 adult male 05day    32 adult male 30day    29 adult female 01day    58 adult female 05day    79 adult female 30day    72 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+    208 Schneider line 2 Sg4    96 embryonic 1182-4H    146 embryonic GM2    0 embryonic Kc167    232 embryonic S1    107 embryonic S3    161 leg disc CME L1    82 wing disc CME-W2    161 wing disc ML-DmD8    157 wing disc ML-DmD9    75 wing disc ML-DmD16-c3    54 wing disc ML-DmD21    199 wing disc ML-DmD32    120 haltere disc ML-DmD17-c3    130 eye-antennal disc ML-DmD11    64 antennal disc ML-DmD20-c5    108 mixed discs ML-DmD4-c1    86 CNS ML-DmBG1-c1    88 CNS ML-DmBG2-c2    107 tumorous blood cells mbn2    214 ovary fGS/OSS    116 ovary OSC    140 ovary OSS    65 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+  (208) Schneider line 2 Sg4  (96) embryonic 1182-4H  (146) embryonic GM2    0 embryonic Kc167  (232) embryonic S1  (107) embryonic S3  (161) leg disc CME L1  (82) wing disc CME-W2  (161) wing disc ML-DmD8  (157) wing disc ML-DmD9  (75) wing disc ML-DmD16-c3  (54) wing disc ML-DmD21  (199) wing disc ML-DmD32  (120) haltere disc ML-DmD17-c3  (130) eye-antennal disc ML-DmD11  (64) antennal disc ML-DmD20-c5  (108) mixed discs ML-DmD4-c1  (86) CNS ML-DmBG1-c1  (88) CNS ML-DmBG2-c2  (107) tumorous blood cells mbn2  (214) ovary fGS/OSS  (116) ovary OSC  (140) ovary OSS  (65) Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Cell Line   Expression Level Schneider line 2 S2R+  (208) Schneider line 2 Sg4    96 embryonic 1182-4H  (146) embryonic GM2    0 embryonic Kc167  (232) embryonic S1  107 embryonic S3  (161) leg disc CME L1    82 wing disc CME-W2  (161) wing disc ML-DmD8  (157) wing disc ML-DmD9    75 wing disc ML-DmD16-c3    54 wing disc ML-DmD21  (199) wing disc ML-DmD32  (120) haltere disc ML-DmD17-c3  (130) eye-antennal disc ML-DmD11    64 antennal disc ML-DmD20-c5  (108) mixed discs ML-DmD4-c1    86 CNS ML-DmBG1-c1    88 CNS ML-DmBG2-c2  107 tumorous blood cells mbn2  (214) ovary fGS/OSS  (116) ovary OSC  (140) ovary OSS    65 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+    208 Schneider line 2 Sg4    96 embryonic 1182-4H    146 embryonic GM2    0 embryonic Kc167    232 embryonic S1    107 embryonic S3    161 leg disc CME L1    82 wing disc CME-W2    161 wing disc ML-DmD8    157 wing disc ML-DmD9    75 wing disc ML-DmD16-c3    54 wing disc ML-DmD21    199 wing disc ML-DmD32    120 haltere disc ML-DmD17-c3    130 eye-antennal disc ML-DmD11    64 antennal disc ML-DmD20-c5    108 mixed discs ML-DmD4-c1    86 CNS ML-DmBG1-c1    88 CNS ML-DmBG2-c2    107 tumorous blood cells mbn2    214 ovary fGS/OSS    116 ovary OSC    140 ovary OSS    65 Expression Level Scale  Extremely high  log, scaled to maximum expression level Cell Line   Expression Level Schneider line 2 S2R+    208 Schneider line 2 Sg4    96 embryonic 1182-4H    146 embryonic GM2    0 embryonic Kc167    232 embryonic S1    107 embryonic S3    161 leg disc CME L1    82 wing disc CME-W2    161 wing disc ML-DmD8    157 wing disc ML-DmD9    75 wing disc ML-DmD16-c3    54 wing disc ML-DmD21    199 wing disc ML-DmD32    120 haltere disc ML-DmD17-c3    130 eye-antennal disc ML-DmD11    64 antennal disc ML-DmD20-c5    108 mixed discs ML-DmD4-c1    86 CNS ML-DmBG1-c1    88 CNS ML-DmBG2-c2    107 tumorous blood cells mbn2    214 ovary fGS/OSS    116 ovary OSC    140 ovary OSS    65 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+  (208) Schneider line 2 Sg4  (96) embryonic 1182-4H  (146) embryonic GM2    0 embryonic Kc167  (232) embryonic S1  (107) embryonic S3  (161) leg disc CME L1  (82) wing disc CME-W2  (161) wing disc ML-DmD8  (157) wing disc ML-DmD9  (75) wing disc ML-DmD16-c3  (54) wing disc ML-DmD21  (199) wing disc ML-DmD32  (120) haltere disc ML-DmD17-c3  (130) eye-antennal disc ML-DmD11  (64) antennal disc ML-DmD20-c5  (108) mixed discs ML-DmD4-c1  (86) CNS ML-DmBG1-c1  (88) CNS ML-DmBG2-c2  (107) tumorous blood cells mbn2  (214) ovary fGS/OSS  (116) ovary OSC  (140) ovary OSS  (65) Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Cell Line   Expression Level Schneider line 2 S2R+  (208) Schneider line 2 Sg4    96 embryonic 1182-4H  (146) embryonic GM2    0 embryonic Kc167  (232) embryonic S1  107 embryonic S3  (161) leg disc CME L1    82 wing disc CME-W2  (161) wing disc ML-DmD8  (157) wing disc ML-DmD9    75 wing disc ML-DmD16-c3    54 wing disc ML-DmD21  (199) wing disc ML-DmD32  120 haltere disc ML-DmD17-c3  130 eye-antennal disc ML-DmD11    64 antennal disc ML-DmD20-c5  108 mixed discs ML-DmD4-c1    86 CNS ML-DmBG1-c1    88 CNS ML-DmBG2-c2  107 tumorous blood cells mbn2  (214) ovary fGS/OSS  116 ovary OSC  140 ovary OSS    65 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+    208 Schneider line 2 Sg4    96 embryonic 1182-4H    146 embryonic GM2    0 embryonic Kc167    232 embryonic S1    107 embryonic S3    161 leg disc CME L1    82 wing disc CME-W2    161 wing disc ML-DmD8    157 wing disc ML-DmD9    75 wing disc ML-DmD16-c3    54 wing disc ML-DmD21    199 wing disc ML-DmD32    120 haltere disc ML-DmD17-c3    130 eye-antennal disc ML-DmD11    64 antennal disc ML-DmD20-c5    108 mixed discs ML-DmD4-c1    86 CNS ML-DmBG1-c1    88 CNS ML-DmBG2-c2    107 tumorous blood cells mbn2    214 ovary fGS/OSS    116 ovary OSC    140 ovary OSS    65 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    76 cold shock, 4-day adult    55 heat shock, 4-day adult    80 Cadmium 50 mM 6 hrs, larvae L3    87 Cadmium 50 mM 12 hrs, larvae L3    152 Cadmium 50 mM 48 hrs, 4-day adult    69 Cadmium 100 mM 48 hrs, 4-day adult    78 Copper 0.5 mM 12 hrs, larvae L3    66 Copper 15 mM 48 hrs, 4-day adult    68 Zinc 5 mM 12 hrs, larvae L3    54 Zinc 4.5 mM 48 hrs, 4-day adult    48 Ethanol 2.5% 3 hrs, larvae L3    96 Ethanol 5% 3 hrs, larvae L3    86 Ethanol 10% 3 hrs, larvae L3    76 Caffeine 1.5 mg/ml 4 hrs, larvae L3    83 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    79 Caffeine 25 mg/ml 48 hrs, 4-day adult    51 Paraquat 5 mM 48 hrs, 4-day adult    72 Paraquat 10 mM 48 hrs, 4-day adult    76 Rotenone 2 μg 12 hrs, larvae L3    91 Rotenone 8 μg 12 hrs, larvae L3    73 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Moderate expression Treatment   Expression Level extended cold, 4-day adult  (76) cold shock, 4-day adult  (55) heat shock, 4-day adult  (80) Cadmium 50 mM 6 hrs, larvae L3  (87) Cadmium 50 mM 12 hrs, larvae L3  (152) Cadmium 50 mM 48 hrs, 4-day adult  (69) Cadmium 100 mM 48 hrs, 4-day adult  (78) Copper 0.5 mM 12 hrs, larvae L3  (66) Copper 15 mM 48 hrs, 4-day adult  (68) Zinc 5 mM 12 hrs, larvae L3  (54) Zinc 4.5 mM 48 hrs, 4-day adult  (48) Ethanol 2.5% 3 hrs, larvae L3  (96) Ethanol 5% 3 hrs, larvae L3  (86) Ethanol 10% 3 hrs, larvae L3  (76) Caffeine 1.5 mg/ml 4 hrs, larvae L3  (83) Caffeine 2.5 mg/ml 48 hrs, 4-day adult  (79) Caffeine 25 mg/ml 48 hrs, 4-day adult  (51) Paraquat 5 mM 48 hrs, 4-day adult  (72) Paraquat 10 mM 48 hrs, 4-day adult  (76) Rotenone 2 μg 12 hrs, larvae L3  (91) Rotenone 8 μg 12 hrs, larvae L3  (73) Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Treatment   Expression Level extended cold, 4-day adult    76 cold shock, 4-day adult    55 heat shock, 4-day adult    80 Cadmium 50 mM 6 hrs, larvae L3    87 Cadmium 50 mM 12 hrs, larvae L3  (152) Cadmium 50 mM 48 hrs, 4-day adult    69 Cadmium 100 mM 48 hrs, 4-day adult    78 Copper 0.5 mM 12 hrs, larvae L3    66 Copper 15 mM 48 hrs, 4-day adult    68 Zinc 5 mM 12 hrs, larvae L3    54 Zinc 4.5 mM 48 hrs, 4-day adult    48 Ethanol 2.5% 3 hrs, larvae L3    96 Ethanol 5% 3 hrs, larvae L3    86 Ethanol 10% 3 hrs, larvae L3    76 Caffeine 1.5 mg/ml 4 hrs, larvae L3    83 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    79 Caffeine 25 mg/ml 48 hrs, 4-day adult    51 Paraquat 5 mM 48 hrs, 4-day adult    72 Paraquat 10 mM 48 hrs, 4-day adult    76 Rotenone 2 μg 12 hrs, larvae L3    91 Rotenone 8 μg 12 hrs, larvae L3    73 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    76 cold shock, 4-day adult    55 heat shock, 4-day adult    80 Cadmium 50 mM 6 hrs, larvae L3    87 Cadmium 50 mM 12 hrs, larvae L3    152 Cadmium 50 mM 48 hrs, 4-day adult    69 Cadmium 100 mM 48 hrs, 4-day adult    78 Copper 0.5 mM 12 hrs, larvae L3    66 Copper 15 mM 48 hrs, 4-day adult    68 Zinc 5 mM 12 hrs, larvae L3    54 Zinc 4.5 mM 48 hrs, 4-day adult    48 Ethanol 2.5% 3 hrs, larvae L3    96 Ethanol 5% 3 hrs, larvae L3    86 Ethanol 10% 3 hrs, larvae L3    76 Caffeine 1.5 mg/ml 4 hrs, larvae L3    83 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    79 Caffeine 25 mg/ml 48 hrs, 4-day adult    51 Paraquat 5 mM 48 hrs, 4-day adult    72 Paraquat 10 mM 48 hrs, 4-day adult    76 Rotenone 2 μg 12 hrs, larvae L3    91 Rotenone 8 μg 12 hrs, larvae L3    73 Expression Level Scale  Extremely high  log, scaled to maximum expression level Treatment   Expression Level extended cold, 4-day adult    76 cold shock, 4-day adult    55 heat shock, 4-day adult    80 Cadmium 50 mM 6 hrs, larvae L3    87 Cadmium 50 mM 12 hrs, larvae L3    152 Cadmium 50 mM 48 hrs, 4-day adult    69 Cadmium 100 mM 48 hrs, 4-day adult    78 Copper 0.5 mM 12 hrs, larvae L3    66 Copper 15 mM 48 hrs, 4-day adult    68 Zinc 5 mM 12 hrs, larvae L3    54 Zinc 4.5 mM 48 hrs, 4-day adult    48 Ethanol 2.5% 3 hrs, larvae L3    96 Ethanol 5% 3 hrs, larvae L3    86 Ethanol 10% 3 hrs, larvae L3    76 Caffeine 1.5 mg/ml 4 hrs, larvae L3    83 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    79 Caffeine 25 mg/ml 48 hrs, 4-day adult    51 Paraquat 5 mM 48 hrs, 4-day adult    72 Paraquat 10 mM 48 hrs, 4-day adult    76 Rotenone 2 μg 12 hrs, larvae L3    91 Rotenone 8 μg 12 hrs, larvae L3    73 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  (76) cold shock, 4-day adult  (55) heat shock, 4-day adult  (80) Cadmium 50 mM 6 hrs, larvae L3  (87) Cadmium 50 mM 12 hrs, larvae L3  (152) Cadmium 50 mM 48 hrs, 4-day adult  (69) Cadmium 100 mM 48 hrs, 4-day adult  (78) Copper 0.5 mM 12 hrs, larvae L3  (66) Copper 15 mM 48 hrs, 4-day adult  (68) Zinc 5 mM 12 hrs, larvae L3  (54) Zinc 4.5 mM 48 hrs, 4-day adult  (48) Ethanol 2.5% 3 hrs, larvae L3  (96) Ethanol 5% 3 hrs, larvae L3  (86) Ethanol 10% 3 hrs, larvae L3  (76) Caffeine 1.5 mg/ml 4 hrs, larvae L3  (83) Caffeine 2.5 mg/ml 48 hrs, 4-day adult  (79) Caffeine 25 mg/ml 48 hrs, 4-day adult  (51) Paraquat 5 mM 48 hrs, 4-day adult  (72) Paraquat 10 mM 48 hrs, 4-day adult  (76) Rotenone 2 μg 12 hrs, larvae L3  (91) Rotenone 8 μg 12 hrs, larvae L3  (73) Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Treatment   Expression Level extended cold, 4-day adult    76 cold shock, 4-day adult    55 heat shock, 4-day adult    80 Cadmium 50 mM 6 hrs, larvae L3    87 Cadmium 50 mM 12 hrs, larvae L3  (152) Cadmium 50 mM 48 hrs, 4-day adult    69 Cadmium 100 mM 48 hrs, 4-day adult    78 Copper 0.5 mM 12 hrs, larvae L3    66 Copper 15 mM 48 hrs, 4-day adult    68 Zinc 5 mM 12 hrs, larvae L3    54 Zinc 4.5 mM 48 hrs, 4-day adult    48 Ethanol 2.5% 3 hrs, larvae L3    96 Ethanol 5% 3 hrs, larvae L3    86 Ethanol 10% 3 hrs, larvae L3    76 Caffeine 1.5 mg/ml 4 hrs, larvae L3    83 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    79 Caffeine 25 mg/ml 48 hrs, 4-day adult    51 Paraquat 5 mM 48 hrs, 4-day adult    72 Paraquat 10 mM 48 hrs, 4-day adult    76 Rotenone 2 μg 12 hrs, larvae L3    91 Rotenone 8 μg 12 hrs, larvae L3    73 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    76 cold shock, 4-day adult    55 heat shock, 4-day adult    80 Cadmium 50 mM 6 hrs, larvae L3    87 Cadmium 50 mM 12 hrs, larvae L3    152 Cadmium 50 mM 48 hrs, 4-day adult    69 Cadmium 100 mM 48 hrs, 4-day adult    78 Copper 0.5 mM 12 hrs, larvae L3    66 Copper 15 mM 48 hrs, 4-day adult    68 Zinc 5 mM 12 hrs, larvae L3    54 Zinc 4.5 mM 48 hrs, 4-day adult    48 Ethanol 2.5% 3 hrs, larvae L3    96 Ethanol 5% 3 hrs, larvae L3    86 Ethanol 10% 3 hrs, larvae L3    76 Caffeine 1.5 mg/ml 4 hrs, larvae L3    83 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    79 Caffeine 25 mg/ml 48 hrs, 4-day adult    51 Paraquat 5 mM 48 hrs, 4-day adult    72 Paraquat 10 mM 48 hrs, 4-day adult    76 Rotenone 2 μg 12 hrs, larvae L3    91 Rotenone 8 μg 12 hrs, larvae L3    73 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_23 mE1_20_mRNA_expression_cluster_10
External Data & Images
Linkouts	BDGP expression data • LD24448 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0000173 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG18319-RA
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal | embryonic stage | non-rescuable maternal effect (with Df(1)KA10) ben1 lethal | embryonic stage | non-rescuable maternal effect | recessive ben1 lethal | partially | recessive ben1 lethal | pupal stage | recessive ben1 benP2 partially lethal - majority die | pharate adult stage P15(ii), with Scer\GAL4pnr-MD237 benGD1387 Other Phenotypes Allele behavior defective benunspecified behavior defective | recessive ben1 chemical sensitive | recessive benJC10 grooming behavior defective | recessive ben1 jumping defective | recessive ben1 ben2 ben3 ben4 ben5 ben6 locomotor behavior defective ben1 benP2 locomotor behavior defective | recessive ben1 neuroanatomy defective ben1 neuroanatomy defective | recessive ben1 uncoordinated | recessive ben1 visible, with Scer\GAL4pnr-MD237 benGD1387 wild-type ben+t6 ben+t8 ben+t10 ben+t18 ben+t35 Phenotype manifest in Allele centrosome | non-rescuable maternal effect ben1 centrosome | non-rescuable maternal effect (with Df(1)KA10) ben1 chaeta | supernumerary, with Scer\GAL4pnr-MD237 benGD1387 coxal tergal remotor muscle ben1 benP2 giant fiber neuron ben1 lamina anlage ben1 lateral oblique dorsal muscle ben1 benP2 medulla anlage ben1 mesothoracic extracoxal depressor muscle 66 ben1 benP2 mesothoracic tergum, with Scer\GAL4pnr-MD237 benGD1387 optic cartridge ben1 photoreceptor cell R1 ben1 photoreceptor cell R2 ben1 photoreceptor cell R3 ben1 photoreceptor cell R4 ben1 photoreceptor cell R5 ben1 photoreceptor cell R6 ben1 photoreceptor cell R7 ben1 photoreceptor cell R8 ben1 retina ben1 rhabdomere R1 ben1 rhabdomere R2 ben1 rhabdomere R3 ben1 rhabdomere R4 ben1 rhabdomere R5 ben1 rhabdomere R6 ben1 rhabdomere R7 ben1 rhabdomere R8 ben1 spindle | non-rescuable maternal effect ben1 spindle | non-rescuable maternal effect (with Df(1)KA10) ben1 tergosternal muscle ben1 benP2
Classical Alleles ( 14 )
For All Classical Alleles Show
Allele of ben Class Mutagen Stocks Known lesion ben1 amorphic allele - genetic evidence ethyl methanesulfonate 1 Yes benG963 P-element activity 1 -- benGS1195 1 Yes ben+ 0 -- ben2 ethyl methanesulfonate 0 -- ben3 ethyl methanesulfonate 0 -- ben4 ethyl methanesulfonate 0 -- ben5 ethyl methanesulfonate 0 -- ben6 ethyl methanesulfonate 0 -- benGG01484 0 -- benGS5108 0 -- benJC10 P-element activity 0 Yes benP2 loss of function allele P-element activity 0 -- benunspecified 0 --
Alleles Carried on Transgenic Constructs ( 11 )
For All Alleles Carried on Transgenic Constructs Show
Allele of ben Class Mutagen Stocks Known lesion benGD1387 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes benJF03148 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes ben+t10 in vitro construct - deletion 0 Yes ben+t18 in vitro construct - genomic fragment 0 Yes ben+t35 in vitro construct - genomic fragment 0 Yes ben+t6 in vitro construct - genomic fragment 0 Yes ben+t8 in vitro construct - deletion 0 Yes benC87A.MtnA.T:Zzzz\FLAG in vitro construct - site directed mutagenesis 0 -- bendsRNA.cZa in vitro construct - RNAi 0 Yes benKK101929 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes benMtnA.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(1)AR2 (Muralidhar and Thomas, 1993) Df(1)benCO1 (Oh et al., 1994) Df(1)benCO2 (Oh et al., 1994) Df(1)g (Edgecomb et al., 1991) Df(1)HA92 (Thomas and Wyman, 1984, Edgecomb et al., 1993, Muralidhar and Thomas, 1993, Tanouye et al., 1987, Oh et al., 1994, Edgecomb et al., 1991)
Not disrupted in	Df(1)AR10 (Muralidhar and Thomas, 1993) Df(1)KA9 (Oh et al., 1994) Df(1)RK2 (Muralidhar and Thomas, 1993, Oh et al., 1994) Ts(1Lt;YLt)B166+Ts(1Rt;YLt)B89 (Lindsley and Zimm, 1992)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD1387} NA P{KK101929} NA P{TRiP.JF03148} NA characterization construct P{89a} NA P{ben.NB} NA P{ben+8} NA P{ben+t6} NA P{ben+t18} NA
Insertions
	Type of insertions Name Expression data insertion of enhancer trap P{lacW}l(1)12DaPL25 No P{lacW}l(1)G0186G0186 No P{PZ}benJC10 No insertion of mobile activating element P{EP}benG963 NA P{GSV1}benGS1195 NA P{GSV2}benGS5108 NA P{Mae-UAS.6.11}benGG01484 NA miscellaneous insertions P{}benP2 NA
Gene Ontology: Function, Process & Cellular Component ( 10 unique terms )
Terms Based on Experimental Evidence ( 7 terms )
Molecular Function ( 0 terms)
Biological Process
CV term Evidence References
axonogenesis inferred from mutant phenotype (Thomas and Wyman, 1982) axon target recognition inferred from mutant phenotype (Muralidhar and Thomas, 1993) flight behavior inferred from mutant phenotype (Oh et al., 1994) grooming behavior inferred from mutant phenotype (Phillis et al., 1993) jump response inferred from mutant phenotype (Thomas and Wyman, 1982) photoreceptor cell morphogenesis inferred from mutant phenotype (Muralidhar and Thomas, 1993) response to anesthetic inferred from mutant phenotype (Campbell et al., 2009)
Cellular Component ( 0 terms)
Terms Based on Predictions or Assertions ( 4 terms )
Molecular Function
CV term Evidence References
acid-amino acid ligase activity inferred from electronic annotation with InterPro:IPR000608 (FlyBase Curators et al., 2004-) ubiquitin-protein ligase activity inferred from sequence or structural similarity (Oh et al., 1994) non-traceable author statement (Swiss-Prot Project Members, 2001.8.20, Muralidhar, 1993.10.1)
Biological Process
CV term Evidence References
grooming behavior traceable author statement (Oh et al., 1994) protein ubiquitination non-traceable author statement (Swiss-Prot Project Members, 2001.8.20)
Cellular Component ( 0 terms)
Sequence Ontology: Class of Gene
	nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
protein-protein
Interacting group Assay References
ben - CG14024 anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) ben - Ent2 anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) ben - Fatp anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) ben - Rab18 anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) ben - Rab5 anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) ben - Rheb anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) ben - Tsp29Fb anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) ben - Uba1 anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based (Mintseris et al., 2009.11.23, Guruharsha et al., 2011)
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions ben allele Gene References
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 58728 DroID - A comprehensive database of gene and protein interactions. • FBgn0000173 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 32358 SignaLink - A protein-protein and protein-miRNA interaction database with multi-layered structure and multiple confidence scores. • P35128
Orthologs
OrthoDB Orthologs (47) - 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 EOG68WCNM Dipteran inclusive ortholog search EOG6N30Z8 Insect inclusive ortholog search EOG6FQZP4 Arthropod inclusive ortholog search EOG6VMCXT Metazoa inclusive ortholog search EOG6V1KVV
Orthologs in Drosophila Species (EOG68WCNM)
Organism Common Name Gene AAA Syntenic Ortholog Multiple Dmel Genes in this Orthologous Group Drosophila melanogaster fruit fly  Dmel\ben     Drosophila simulans   Dsim\GD17149 Y   Drosophila sechellia   Dsec\GM17649 Y   Drosophila erecta   Dere\GG17819 Y   Drosophila yakuba   Dyak\GE17114 Y   Drosophila ananassae   Dana\GF19231 Y   Drosophila pseudoobscura pseudoobscura   Dpse\GA14886 Y   Drosophila persimilis   Dper\GL14825 Y   Drosophila willistoni   Dwil\GK25846 Y   Drosophila virilis   Dvir\GJ23435     Drosophila virilis   Dvir\GJ19035 Y   Drosophila mojavensis   Dmoj\GI22711     Drosophila mojavensis   Dmoj\GI21493 Y   Drosophila grimshawi   Dgri\GH22592     Drosophila grimshawi   Dgri\GH17677
Orthologs in non-Drosophila Dipterans (EOG6N30Z8)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Aedes aegypti Yellow fever mosquito  Aaeg\AAEL014642 Y Aedes aegypti Yellow fever mosquito  Aaeg\AAEL014639 Y Anopheles gambiae Malaria mosquito  Agam\AGAP005197 Y Culex quinquefasciatus Southern house mosquito  Cqui\CPIJ006567 Y
Orthologs in non-Dipteran Insects (EOG6FQZP4)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Apis mellifera Western honey bee  Amel\Ubc13 Y Nasonia vitripennis Parasitic wasp  Nvit\Nasvi2EG011477 Y Acromyrmex echinatior Panamanian leafcutter ant  Aech\AECH18999 Y Atta cephalotes Leafcutter ant  Acep\ACEP27525 Y Camponotus floridanus Florida carpenter ant  Cflo\CFLO18469 Y Harpegnathos saltator Jerdons jumping ant  Hsal\HSAL16636 Y Linepithema humile Argentine ant  Lhum\LH21961 Y Pogonomyrmex barbatus Red harvester ant  Pbar\PB21899 Y Solenopsis invicta Red fire ant  Sinv\SINV16216 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI006341 Y Bombyx mori Silkmoth  Bmor\BGIBMGA002559 Y Pediculus humanus Human body louse  Phum\PHUM593530 Y Tribolium castaneum Red flour beetle  Tcas\TC001755 Y
Orthologs in non-Insect Arthropods (EOG6VMCXT)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Daphnia pulex Water flea  Dpux\G3pdh Y Ixodes scapularis Deer tick  Isca\ISCW006743 Y
Orthologs in non-Arthropod Metazoa (EOG6V1KVV)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Caenorhabditis elegans Nematode  Cele\ubc-13 Y Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_018598gn Y Danio rerio Zebrafish  Drer\ube2nl Y Danio rerio Zebrafish  Drer\ube2n Y Xenopus tropicalis Western clawed frog  Xtro\ube2n Y Gallus gallus Domestic chicken  Ggal\UBE2N Y Mus musculus House mouse  Mmus\Ube2nl Y Mus musculus House mouse  Mmus\Ube2n Y Rattus norvegicus Norway rat  Rnor\ENSRNOG00000017935 Y Rattus norvegicus Norway rat  Rnor\Ube2n Y Rattus norvegicus Norway rat  Rnor\ENSRNOG00000039638 Y Homo sapiens Human  Hsap\UBE2N Y Homo sapiens Human  Hsap\UBE2NL Y
Human Orthologs (2)
Gene OMIM HGNC Hsap\UBE2N     Hsap\UBE2NL
AAA Orthologs (10) based on analysis using Dmel annotation version 4.3
Organism Gene Drosophila simulans Dsim\GD17149 Drosophila sechellia Dsec\GM17649 Drosophila erecta Dere\GG17819 Drosophila yakuba Dyak\GE17114 Drosophila ananassae Dana\GF19231 Drosophila pseudoobscura pseudoobscura Dpse\GA14886 Drosophila persimilis Dper\GL14825 Drosophila willistoni Dwil\GK25846 Drosophila virilis Dvir\GJ19035 Drosophila mojavensis Dmoj\GI21493
Stocks & Reagents
Stocks Listed in FlyBase ( 5 )
Bloomington	33464 w* P{EP}benG963 28721 y1 v1; P{TRiP.JF03148}attP2 9565 w118 ben1/C(1)A, y1
Kyoto	205033 y1 w67c23 P{GSV1}benGS1195/Binsinscy
VDRC	v9413 w1118; P{GD1387}v9413
Genomic Clones ( 1 )
	BACR22M24 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 118 )
	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	LD24448
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	GH04055 GH08861 GH10458 GH10956 GH22610 GM02759 GM30629 LD14534 LD16328 LD31794 LD32667 RE13202 RE33233 RE34174 RE35152 RE35823 RE40629 RE48261 RE52552 RE63604 RE65493 RH40204
Other clones	5148 10976 19380 20092 30213 34591 36563 36732 40192 42221 56608 71284 72010 72581 79647 83979 101538 102360 107465 108372 108635 120468 165149 183973 193435 214289.12 214511.10 214573.6 273279 277387 277529 283384 562235 CK01.11A.E7 CK01.15B.C10 CK01.26B.H4 CK01.63A.F7 CK01.64B.D12 CK01.87A.C6 CK01.97C.G6 CK01.101C.C11 CK01.118A.A12 CK01.129D.G11 CK01.170B.D8 Dmel_neb_012845_0077_1436 EC051299 EK012920 EK052336 EK065146 EK067607 EK075559 EK077912 EK085709 EK090727 EK096226 EK103009 EK104448 EK111039 EK123962 EK132360 EK133812 EK144846 EK147110 EK147523 EK153918 EK157341 EK169022 EK198927 EK200349 EK201515 EK209858 EK218463 EK227941 EK240657 EK270612 EK296733 EK304823 EK304856 EK308837 EN13947 EP09458 EP10043 EP19248 ESG015b.E21_C12 ESG016b.E21_C12 ESG017b.E21_B12 FGA001C11 FGA001D02 FGH112C08 FGM216B02 IP05365 IP05377 IP05565 RH08946 UUGC0572
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0000173 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 32358
Antibody Information
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of	Source for identity of: ben CG18319 (Zheng, 1999.11)
Source for database merge of	Source for merge of: ben anon-WO03040301.242 (FlyBase, 1996-)
Additional comments	"l(1)G0186" may affect "ben". (Peter et al., 2002) Source for merge of ben anon-WO03040301.242 was sequence comparison (date:051113). (FlyBase, 1996-)
Other Comments
	dsRNA made from templates generated with primers directed against this gene. (Mukai et al., 2007) 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) ben mRNA is terminated at the poly(A) signal provided by the Ste intergenic spacer 3' untranslated region which is present upstream of the neighbouring Ste12DOR coding region. Overall orientation not stated: ben+ Ste12DOR+ (Gvozdev et al., 2000) Mutants have pleiotropic phenotypes affecting thoracic muscle patterning, pupal mortality and post-eclosion mortality. (Edgecomb et al., 1993) Mutations in ben cause a disruption in the recognition event leading to synapse formation between the giant fibre (GF) and the tergotrochanteral jump muscle motorneuron (TTMmn) giving rise to the GF phenotype. Also mutants exhibit morphological abnormalities within the visual system. Results suggest a role for ben for ubiquitin-mediated protein modification in nervous system development, including the regulation of synaptic connectivity. (Muralidhar and Thomas, 1993) Adults have an aberrant startle response; they do not jump when presented with a lights-off stimulus. The cervical giant fiber, a brain neuron, has abnormal morphology. The normal giant fiber terminates in the thorax at a synapse onto the motoneuron innervating the tergo-trochanteral (TT) muscle (= jump muscle). In ben, this synapse is abnormal or absent. The normal lateral bend of the giant fiber toward the TT motoneuron in the mesothoracic neuromere is absent; the axon usually terminates at the midline with fine branches extending from its tip. Latency of response of the TT muscle to stimulation of the giant fiber is abnormally long, and muscles cannot follow stimulation at rates above 5 Hz. Rhabdomeres of the photoreceptor cells in the ommatidia heart shaped in cross-section with indentation centrally oriented rather than round as is normal. Furthermore, the axons from photoreceptor cells R7 and R8 fail to make the right-angle turn into the optic medulla after traversing the lamina. ben flies choose visible over ultraviolet wave lengths whereas wild-type flies make the opposite choice. (Lindsley and Zimm, 1992) Less than 25% ben flies show abnormalities in the dorsal attachment of the tergotrochantral muscle. (Edgecomb et al., 1991) Behavioural data suggests antennal and maxillary basiconic sensilla may be important receptors for short chain alcohols and organic acids but less crucial for acetates, aldehydes and ketones. (Venard and Stocker, 1991) Mutations have a defect in the pathway between the giant fibre and jump muscle, the giant fiber fails to make a bend which would normally bring it into proximity with the jump muscle motoneuron. (Tanouye et al., 1987) Mutations interrupt one neural connection in the giant fibre pathway that mediates the jump/flight escape response. (Tanouye et al., 1986) Mutations disrupt the synaptic transmission between the giant fibre (GF) system and the tergotrochanteral muscle (TTM, jumping muscle) motor neuron. (Thomas and Wyman, 1984) Mutation in ben deletes one of the processes of the giant fibre leaving the other portions functionally unaltered. (Thomas and Wyman, 1982)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_078595 NM_001169281 NM_001258734 NM_001272611 NM_001272612 RefSeq (Proteins) • NP_511150 NP_001162752 NP_001245663 NP_001259540 NP_001259541 DDBJ / EMBL / GenBank • AA820626 AAA28392 AAB30753 AAF48338 AAL39672 ACZ95287 AF192309 AFH07377 AGB95382 AGB95383 AI455478 AX777388 AY069527 L20126 S70118 Entrez Gene - A searchable database of RefSeq genes. • 32358 UniProtKB/Swiss-Prot • P35128 UniProtKB/TrEMBL • E1JJN1
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Ubiquitin-conjugating enzyme, E2 (IPR000608) Ubiquitin-conjugating enzyme/RWD-like (IPR016135) Ubiquitin-conjugating enzyme, active site (IPR023313) BDGP expression data - Patterns of gene expression in Drosophila embryogenesis • LD24448
Linkouts
	BioGRID - A database of protein and genetic interactions • 58728 DroID - A comprehensive database of gene and protein interactions. • FBgn0000173 DRSC - Results from RNAi screens. • FBgn0000173 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0000173 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG18319-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0000173 FlyReactome - A curated knowledgebase of Drosophila melanogaster pathways • 104205 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 32358 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 32358 modMine - Data generated by the modENCODE project. • FBgn0000173 SignaLink - A protein-protein and protein-miRNA interaction database with multi-layered structure and multiple confidence scores. • P35128
Synonyms & Secondary IDs ( 13 )
Reported As
Symbol Synonym	anon-WO03040301.242   Ben (Zhou et al., 2002, Merkle et al., 2007, Ying et al., 2011) ben (Laviolette et al., 2005, Garcia et al., 2007, Mukai et al., 2007, Hill-Burns and Clark, 2010, Uthaman et al., 2008, Merkle et al., 2009, Campbell et al., 2009, Franco et al., 2011) BEN (Merkle and Lee, 2008, Merkle et al., 2009) Bend (Paquette et al., 2010) CG18319 (Laviolette et al., 2005, Toledano-Katchalski et al., 2007, Perkins et al., 2009, Keleman et al., 2009.8.5, Cammarato et al., 2011, Ying et al., 2011) nj-262   ubc13 (Silverman et al., 2008) Ubc13 (Herman-Bachinsky et al., 2007, Cronin et al., 2009, Paquette et al., 2010) UbcD3 (Matuschewski et al., 1996, Michelle et al., 2009)
Name Synonym	bendless (Muralidhar, 1993.10.1, Zhou et al., 2005, Silverman et al., 2008, Murphey and Godenschwege, 2002, Uthaman et al., 2008, Paquette et al., 2010, Cammarato et al., 2011) Bendless (Meinander et al., 2012, Merkle et al., 2009) nonjumper-262
Secondary FlyBase IDs
	FBgn0066817
References ( 88 )
Generate a list of	All references relating to this gene ( 88 )
List References by type	Research paper  ( 33 ) Supplementary material  ( 10 ) Review  ( 7 ) Personal communication to FlyBase  ( 8 ) Abstract  ( 17 ) FlyBase analysis  ( 3 ) DNA/RNA sequence record  ( 2 ) Conference report  ( 1 ) Curated genome annotation  ( 3 ) Book  ( 1 ) Protein sequence record  ( 1 ) Website  ( 1 ) Patent  ( 1 )
Recent research papers ( 4 )
	Ma et al., 2012, Cell Res. 22(2): 425--431 NOPO modulates Egr-induced JNK-independent cell death in Drosophila. [FBrf0217390] Meinander et al., 2012, EMBO J. 31(12): 2770--2783 Ubiquitylation of the initiator caspase DREDD is required for innate immune signalling. [FBrf0218688] Franco et al., 2011, Mol. Cell. Proteomics 10(5): M110.002188 A Novel Strategy to Isolate Ubiquitin Conjugates Reveals Wide Role for Ubiquitination during Neural Development. [FBrf0213919] Ying et al., 2011, PLoS ONE 6(9): e23863 Comprehensively Surveying Structure and Function of RING Domains from Drosophila melanogaster. [FBrf0215242] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2011 Show reviews ...