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Gene Dmel\piwi

General Information
Symbol	Dmel\piwi	Species	D. melanogaster
Name	piwi	Annotation symbol	CG6122
Feature type	protein_coding_gene	FlyBase ID	FBgn0004872
Gene Model Status	Current	Stock availability	11 publicly available
Genomic Location
Chromosome (arm)	2L	Recombination map
Cytogenetic map	32C1-32C1	Sequence location	2L:10,982,205..10,987,420 [-]
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 piwi is referred to in FlyBase by the symbol Dmel\piwi (CG6122, FBgn0004872). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: protein binding; RNA binding. There is experimental evidence for 11 unique biological process terms, many of which group under: biological regulation; single-organism developmental process; multicellular organism reproduction; stem cell division; gamete generation; gene silencing; negative regulation of cellular biosynthetic process; cellular component organization or biogenesis; male germ-line stem cell division; pole cell development; negative regulation of DNA recombination; regulation of developmental process; positive regulation of nuclear-transcribed mRNA poly(A) tail shortening; negative regulation of metabolic process. 36 alleles are reported. The phenotypes of these alleles are annotated with: multicellular structure; organ system; anatomical structure; ovariole; multi-tissue structure; material anatomical entity; female organism; cytoplasmic part; acellular anatomical structure; organelle; portion of tissue. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: Argonaute/Dicer protein, PAZ; Ribonuclease H-like domain; Stem cell self-renewal protein Piwi. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of high expression to a trough of very low expression. Peak expression observed within 00-06 hour embryonic stages, in adult female stages. Summary of FlyAtlas Anatomical Expression Data: Expression at high levels in the following post-embryonic organs or tissues: adult ovary. Comments on Affy2 ProbeSet: ProbeSet 1641305_at completely aligns to an exonic region of the only FlyBase-annotated transcript isoform of piwi. Gene sequence location is 2L:10982205..10987420.
User Contributed Data
	Beta version, feedback welcome
External Summaries
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 heterochromatin organization involved in chromatin silencing
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 32C1-32C1   Limits computationally determined from genome sequence between P{lacW}l(2)k05812k05812&P{lacW}l(2)k09104k09104 and P{EP}ppoEP2478&P{lacW}l(2)k15817k15817
Experimentally Determined Cytological Location
Cytogenetic map Notes References 32B-32C   (determined by in situ hybridisation)   (Spradling et al., 1999) 32C-32C   (determined by in situ hybridisation)   (Lin and Spradling, 1997)
Experimentally Determined Recombination Data
Location
Left of (cM)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\piwi 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 5.50 Low-frequency RNA-Seq exon junction(s) not annotated.
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) piwi-RA FBtr0080166  NM_057527   3073   843 piwi-RB FBtr0340227     2969   843
Additional Transcript Data & Comments
Reported size (kB)	3.1 (northern blot) (Cox et al., 1998)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein piwi-PA   FBpp0079755   97.2   843   10.00     NP_476875   AAF53043 piwi-PB   FBpp0309202   97.2   843   10.00
Additional Polypeptide Data & Comments
Reported size (kDa)	843 (aa); 97.2 (kD predicted) (Cox et al., 1998)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Argonaute/Dicer protein, PAZ (IPR003100) Stem cell self-renewal protein Piwi (IPR003165) Ribonuclease H-like domain (IPR012337)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA202789   AA802575   AA978826   AE014134 AAF53043   AC092226   AF104354 AAD08704   AF104355 AAD08705   AI061964   AQ025014   AW941950   BG637778   BG637781   BI163833   BI168498   BI214891   BI241150   BI354756   BI372643   BP542472   BP542762   BP543149   BP543150   BP543973   BP543974   BP553307   BT011097   BT011138   BT071791   CO177475   CO179730   CO186326   CO324805   CZ478664   EC057157   EC063398   EC207389   EC241555   EC255169   EV580069   GH753605   GH771553   GH773386   GH774539   GH782767   GH783947   GH784070   GH786807   GH789252   GH798223   GH802358   GH808269   GH809208   GH814120   GH814491   GH815147   GH816164   GH822098   GH826030   GH830182   GH835937   GH838619   GH841491   GH841794   GH849191   GH853553   GH861045   GH865154   GH868393   GH869010   GH869015   GH869374   GH872701   GH872834   GH882114   GH883780   GH884871   GH895879   GH898664   GH899792   GH900445   GH903416   GH905448   GH906054   GH909491   GH916595   GH922257   GH922543   GH923727   GH923815   GH926241   GH933216   GH937264   GH939474   GH940007
UniProtKB/Swiss-Prot	Q9VKM1
UniProtKB/TrEMBL
Mapped Features
Mapped Features have been reorganized, please see this article for details. Additional mapped features and mutations can be found on GBrowse or related reports. Type Symbol & Location Additional Notes References
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage | early     (Cox et al., 1998) embryonic stage 1 -- 3   pole plasm   (Fisher et al., 2012) Comment:maternally deposited organism   • pole plasm (Fisher et al., 2012) Comment:maternally deposited embryonic stage 4 -- 6   organism   (Fisher et al., 2012) Comment:rapidly degraded organism | faint & ubiquitous   (Fisher et al., 2012) pole cell   (Fisher et al., 2012) embryonic stage 7 -- 8   pole cell   (Fisher et al., 2012) embryonic stage 9 -- 10   germline cell   (Fisher et al., 2012) embryonic stage 11 -- 12   germline cell   (Fisher et al., 2012) embryonic stage 13 -- 16   germline cell   (Fisher et al., 2012) gonad   (Fisher et al., 2012) adult stage & oogenesis   germarium region 2b   (Cox et al., 1998) germarium region 2a   (Cox et al., 1998) epithelial sheath   (Cox et al., 1998) ovary   (Cox et al., 1998) terminal filament basal cell   (Cox et al., 1998)
Additional Descriptive Data
	piwi transcript is detected in germline and soma cells during oogenesis. In the germarium, expression is detected in the terminal filament cells and the epithelial sheath cells. In germ cells, expression is detected as early as germarium region 2, continues during stages S1-S6 of oogenesis, is much diminished but still detectable between S7-S9, and is again at high levels at stage S10. In early embryos, expression is uniform. (Cox et al., 1998)
Marker for
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
Subcellular Localization (GO Cellular Component)
CV term Evidence References
chromocenter inferred from direct assay (Brower-Toland et al., 2007) cytoplasm inferred from direct assay (Zamparini et al., 2011) euchromatin inferred from direct assay (Brower-Toland et al., 2007) nucleoplasm inferred from direct assay (Lau et al., 2009) inferred from direct assay (assigned by UniProtKB) (Cox et al., 2000) nucleus inferred from direct assay (Brennecke et al., 2007, Saito et al., 2006, Grimaud et al., 2006, Zamparini et al., 2011, Olivieri et al., 2012) polytene chromosome inferred from direct assay (Brower-Toland et al., 2007) telomeric heterochromatin inferred from direct assay (Brower-Toland et al., 2007) Yb body inferred from physical interaction with armi (Handler et al., 2011)
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: Expression at high levels in the following post-embryonic organs or tissues: adult ovary. [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    11.875 Larval Midgut    9 Larval Hindgut    9.3 Larval Malpighian Tubules    9.3 Larval Fat Body    49.8 Larval Salivary Gland    6.9 Larval Trachea    5.35 Larval Carcass    8.45 Adult Head    6.3 Adult Eye    5.375 Adult Brain    10.2 Adult Thoracic-Abdominal Ganglion    10.2 Adult Crop    11.1 Adult Midgut    9.2 Adult Hindgut    10.2 Adult Malpighian Tubules    8.4 Adult Fat Body    13.7 Adult Salivary Gland    8.8 Adult Heart    6.65 Adult VirginFemale Spermatheca    6.5 Adult InseminatedFemale Spermatheca    7.2 Adult Ovary    583.7 Adult Testis    18.2 Adult Male Accessory Gland    7 Adult Carcass    15.4 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    11.875 Larval Midgut    9 Larval Hindgut    9.3 Larval Malpighian Tubules    9.3 Larval Fat Body    49.8 Larval Salivary Gland    6.9 Larval Trachea    5.35 Larval Carcass    8.45 Adult Head    6.3 Adult Eye    5.375 Adult Brain    10.2 Adult Thoracic-Abdominal Ganglion    10.2 Adult Crop    11.1 Adult Midgut    9.2 Adult Hindgut    10.2 Adult Malpighian Tubules    8.4 Adult Fat Body    13.7 Adult Salivary Gland    8.8 Adult Heart    6.65 Adult VirginFemale Spermatheca    6.5 Adult InseminatedFemale Spermatheca    7.2 Adult Ovary  (583.7) Adult Testis    18.2 Adult Male Accessory Gland    7 Adult Carcass    15.4 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    11.875 Larval Midgut    9 Larval Hindgut    9.3 Larval Malpighian Tubules    9.3 Larval Fat Body    49.8 Larval Salivary Gland    6.9 Larval Trachea    5.35 Larval Carcass    8.45 Adult Head    6.3 Adult Eye    5.375 Adult Brain    10.2 Adult Thoracic-Abdominal Ganglion    10.2 Adult Crop    11.1 Adult Midgut    9.2 Adult Hindgut    10.2 Adult Malpighian Tubules    8.4 Adult Fat Body    13.7 Adult Salivary Gland    8.8 Adult Heart    6.65 Adult VirginFemale Spermatheca    6.5 Adult InseminatedFemale Spermatheca    7.2 Adult Ovary    583.7 Adult Testis    18.2 Adult Male Accessory Gland    7 Adult Carcass    15.4 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    11.875 Larval Midgut    9 Larval Hindgut    9.3 Larval Malpighian Tubules    9.3 Larval Fat Body    49.8 Larval Salivary Gland    6.9 Larval Trachea    5.35 Larval Carcass    8.45 Adult Head    6.3 Adult Eye    5.375 Adult Brain    10.2 Adult Thoracic-Abdominal Ganglion    10.2 Adult Crop    11.1 Adult Midgut    9.2 Adult Hindgut    10.2 Adult Malpighian Tubules    8.4 Adult Fat Body    13.7 Adult Salivary Gland    8.8 Adult Heart    6.65 Adult VirginFemale Spermatheca    6.5 Adult InseminatedFemale Spermatheca    7.2 Adult Ovary    583.7 Adult Testis    18.2 Adult Male Accessory Gland    7 Adult Carcass    15.4 Expression Level Scale  Very high  log, scaled to maximum expression level Tissue   Expression Level Larval Central Nervous System    11.875 Larval Midgut    9 Larval Hindgut    9.3 Larval Malpighian Tubules    9.3 Larval Fat Body    49.8 Larval Salivary Gland    6.9 Larval Trachea    5.35 Larval Carcass    8.45 Adult Head    6.3 Adult Eye    5.375 Adult Brain    10.2 Adult Thoracic-Abdominal Ganglion    10.2 Adult Crop    11.1 Adult Midgut    9.2 Adult Hindgut    10.2 Adult Malpighian Tubules    8.4 Adult Fat Body    13.7 Adult Salivary Gland    8.8 Adult Heart    6.65 Adult VirginFemale Spermatheca    6.5 Adult InseminatedFemale Spermatheca    7.2 Adult Ovary    583.7 Adult Testis    18.2 Adult Male Accessory Gland    7 Adult Carcass    15.4 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    11.875 Larval Midgut    9 Larval Hindgut    9.3 Larval Malpighian Tubules    9.3 Larval Fat Body    49.8 Larval Salivary Gland    6.9 Larval Trachea    5.35 Larval Carcass    8.45 Adult Head    6.3 Adult Eye    5.375 Adult Brain    10.2 Adult Thoracic-Abdominal Ganglion    10.2 Adult Crop    11.1 Adult Midgut    9.2 Adult Hindgut    10.2 Adult Malpighian Tubules    8.4 Adult Fat Body    13.7 Adult Salivary Gland    8.8 Adult Heart    6.65 Adult VirginFemale Spermatheca    6.5 Adult InseminatedFemale Spermatheca    7.2 Adult Ovary  583.7 Adult Testis    18.2 Adult Male Accessory Gland    7 Adult Carcass    15.4 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    11.875 Larval Midgut    9 Larval Hindgut    9.3 Larval Malpighian Tubules    9.3 Larval Fat Body    49.8 Larval Salivary Gland    6.9 Larval Trachea    5.35 Larval Carcass    8.45 Adult Head    6.3 Adult Eye    5.375 Adult Brain    10.2 Adult Thoracic-Abdominal Ganglion    10.2 Adult Crop    11.1 Adult Midgut    9.2 Adult Hindgut    10.2 Adult Malpighian Tubules    8.4 Adult Fat Body    13.7 Adult Salivary Gland    8.8 Adult Heart    6.65 Adult VirginFemale Spermatheca    6.5 Adult InseminatedFemale Spermatheca    7.2 Adult Ovary    583.7 Adult Testis    18.2 Adult Male Accessory Gland    7 Adult Carcass    15.4 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    11.875 Larval Midgut    9 Larval Hindgut    9.3 Larval Malpighian Tubules    9.3 Larval Fat Body    49.8 Larval Salivary Gland    6.9 Larval Trachea    5.35 Larval Carcass    8.45 Adult Head    6.3 Adult Eye    5.375 Adult Brain    10.2 Adult Thoracic-Abdominal Ganglion    10.2 Adult Crop    11.1 Adult Midgut    9.2 Adult Hindgut    10.2 Adult Malpighian Tubules    8.4 Adult Fat Body    13.7 Adult Salivary Gland    8.8 Adult Heart    6.65 Adult VirginFemale Spermatheca    6.5 Adult InseminatedFemale Spermatheca    7.2 Adult Ovary    583.7 Adult Testis    18.2 Adult Male Accessory Gland    7 Adult Carcass    15.4 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    6.3   NA  Eye    5.375   NA  Brain    no informative data   11.875  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    no informative data   NA  Crop    no informative data   9  Midgut    9.2   9.3  Hindgut    no informative data   9.3  Malpighian Tubules    8.4   49.8  Fat Body    no informative data   6.9  Salivary Gland    8.8   NA  Heart    6.65   5.35  Trachea    NA   NA  VirginFemale Spermatheca    6.5   NA  InseminatedFemale Spermatheca    7.2   NA  Ovary    583.7   NA  Testis    18.2   NA  Male Accessory Gland    7   8.45  Carcass    no informative data 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    3 central nervous system, larvae L3    0 central nervous system, pupae P8    0 head, virgin 1-day female    0 head, virgin 4-day female    1 head, virgin 20-day female    0 head, mated 1-day female    2 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    1 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    0 digestive system, larvae L3 wandering    0 digestive system, 1-day adult    1 digestive system, 4-day adult    8 digestive system, 20-day adult    5 fat body, larvae L3 wandering    1 fat body, white prepupae    6 fat body, pupae P8    0 carcass, larvae L3 wandering    0 carcass, 1-day adult    3 carcass, 4-day adult    8 carcass, 20-day adult    6 ovary, virgin 4-day female    231 ovary, mated 4-day female    167 testis, mated 4-day male    2 accessory gland, mated 4-day male    0 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    3 central nervous system, larvae L3    0 central nervous system, pupae P8    0 head, virgin 1-day female    0 head, virgin 4-day female    1 head, virgin 20-day female    0 head, mated 1-day female    2 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    1 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    0 digestive system, larvae L3 wandering    0 digestive system, 1-day adult    1 digestive system, 4-day adult    8 digestive system, 20-day adult    5 fat body, larvae L3 wandering    1 fat body, white prepupae    6 fat body, pupae P8    0 carcass, larvae L3 wandering    0 carcass, 1-day adult    3 carcass, 4-day adult    8 carcass, 20-day adult    6 ovary, virgin 4-day female  (231) ovary, mated 4-day female  (167) testis, mated 4-day male    2 accessory gland, mated 4-day male    0 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Tissue   Expression Level imaginal disc, larvae L3 wandering    3 central nervous system, larvae L3    0 central nervous system, pupae P8    0 head, virgin 1-day female    0 head, virgin 4-day female    1 head, virgin 20-day female    0 head, mated 1-day female    2 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    1 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    0 digestive system, larvae L3 wandering    0 digestive system, 1-day adult    1 digestive system, 4-day adult    8 digestive system, 20-day adult    5 fat body, larvae L3 wandering    1 fat body, white prepupae    6 fat body, pupae P8    0 carcass, larvae L3 wandering    0 carcass, 1-day adult    3 carcass, 4-day adult    8 carcass, 20-day adult    6 ovary, virgin 4-day female  (231) ovary, mated 4-day female  (167) testis, mated 4-day male    2 accessory gland, mated 4-day male    0 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    3 central nervous system, larvae L3    0 central nervous system, pupae P8    0 head, virgin 1-day female    0 head, virgin 4-day female    1 head, virgin 20-day female    0 head, mated 1-day female    2 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    1 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    0 digestive system, larvae L3 wandering    0 digestive system, 1-day adult    1 digestive system, 4-day adult    8 digestive system, 20-day adult    5 fat body, larvae L3 wandering    1 fat body, white prepupae    6 fat body, pupae P8    0 carcass, larvae L3 wandering    0 carcass, 1-day adult    3 carcass, 4-day adult    8 carcass, 20-day adult    6 ovary, virgin 4-day female    231 ovary, mated 4-day female    167 testis, mated 4-day male    2 accessory gland, mated 4-day male    0 Expression Level Scale  Extremely high  log, scaled to maximum expression level Tissue   Expression Level imaginal disc, larvae L3 wandering    3 central nervous system, larvae L3    0 central nervous system, pupae P8    0 head, virgin 1-day female    0 head, virgin 4-day female    1 head, virgin 20-day female    0 head, mated 1-day female    2 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    1 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    0 digestive system, larvae L3 wandering    0 digestive system, 1-day adult    1 digestive system, 4-day adult    8 digestive system, 20-day adult    5 fat body, larvae L3 wandering    1 fat body, white prepupae    6 fat body, pupae P8    0 carcass, larvae L3 wandering    0 carcass, 1-day adult    3 carcass, 4-day adult    8 carcass, 20-day adult    6 ovary, virgin 4-day female    231 ovary, mated 4-day female    167 testis, mated 4-day male    2 accessory gland, mated 4-day male    0 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    3 central nervous system, larvae L3    0 central nervous system, pupae P8    0 head, virgin 1-day female    0 head, virgin 4-day female    1 head, virgin 20-day female    0 head, mated 1-day female    2 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    1 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    0 digestive system, larvae L3 wandering    0 digestive system, 1-day adult    1 digestive system, 4-day adult    8 digestive system, 20-day adult    5 fat body, larvae L3 wandering    1 fat body, white prepupae    6 fat body, pupae P8    0 carcass, larvae L3 wandering    0 carcass, 1-day adult    3 carcass, 4-day adult    8 carcass, 20-day adult    6 ovary, virgin 4-day female  (231) ovary, mated 4-day female  (167) testis, mated 4-day male    2 accessory gland, mated 4-day male    0 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Tissue   Expression Level imaginal disc, larvae L3 wandering    3 central nervous system, larvae L3    0 central nervous system, pupae P8    0 head, virgin 1-day female    0 head, virgin 4-day female    1 head, virgin 20-day female    0 head, mated 1-day female    2 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    1 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    0 digestive system, larvae L3 wandering    0 digestive system, 1-day adult    1 digestive system, 4-day adult    8 digestive system, 20-day adult    5 fat body, larvae L3 wandering    1 fat body, white prepupae    6 fat body, pupae P8    0 carcass, larvae L3 wandering    0 carcass, 1-day adult    3 carcass, 4-day adult    8 carcass, 20-day adult    6 ovary, virgin 4-day female  (231) ovary, mated 4-day female  (167) testis, mated 4-day male    2 accessory gland, mated 4-day male    0 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    3 central nervous system, larvae L3    0 central nervous system, pupae P8    0 head, virgin 1-day female    0 head, virgin 4-day female    1 head, virgin 20-day female    0 head, mated 1-day female    2 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    1 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    0 digestive system, larvae L3 wandering    0 digestive system, 1-day adult    1 digestive system, 4-day adult    8 digestive system, 20-day adult    5 fat body, larvae L3 wandering    1 fat body, white prepupae    6 fat body, pupae P8    0 carcass, larvae L3 wandering    0 carcass, 1-day adult    3 carcass, 4-day adult    8 carcass, 20-day adult    6 ovary, virgin 4-day female    231 ovary, mated 4-day female    167 testis, mated 4-day male    2 accessory gland, mated 4-day male    0 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Tissue   Expression Level imaginal disc, larvae L3 wandering     central nervous system, larvae L3     central nervous system, pupae P8     head, virgin 1-day female     head, virgin 4-day female     head, virgin 20-day female     head, mated 1-day female     head, mated 4-day female     head, mated 20-day female     head, mated 1-day male     head, mated 4-day male     head, mated 20-day male     salivary gland, larvae L3 wandering     salivary gland, white prepupae     digestive system, larvae L3 wandering     digestive system, 1-day adult     digestive system, 4-day adult     digestive system, 20-day adult     fat body, larvae L3 wandering     fat body, white prepupae     fat body, pupae P8     carcass, larvae L3 wandering     carcass, 1-day adult     carcass, 4-day adult     carcass, 20-day adult     ovary, virgin 4-day female     ovary, mated 4-day female     testis, mated 4-day male     accessory gland, mated 4-day male
modENCODE Development RNA-Seq
modENCODE Temporal Expression Data (modENCODE_mRNA-Seq_U)    Styles Linear Logarithmic Heatmap    Scales gene max expression Moderate expression bin max High expression bin max Extremely high expression bin max Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of high expression to a trough of very low expression. Peak expression observed within 00-06 hour embryonic stages, in adult female 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    76 embryo 02-04hr    9 embryo 04-06hr    5 embryo 06-08hr    2 embryo 08-10hr    2 embryo 10-12hr    1 embryo 12-14hr    2 embryo 14-16hr    2 embryo 16-18hr    3 embryo 18-20hr    2 embryo 20-22hr    4 embryo 22-24hr    5 larva L1    2 larva L2    2 larva L3 12hr old    1 larva L3 puffstage 1-2    0 larva L3 puffstage 3-6    1 larva L3 puffstage 7-9    1 white prepupae new    1 white prepupae 12hr    2 white prepupae 24hr    2 pupae 2d postWPP    2 pupae 3d postWPP    3 pupae 4d postWPP    7 adult male 01day    1 adult male 05day    1 adult male 30day    1 adult female 01day    37 adult female 05day    76 adult female 30day    69 Expression Level Scale  Very low   Low   Moderate   Moderately high   High  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (76) embryo 02-04hr    9 embryo 04-06hr    5 embryo 06-08hr    2 embryo 08-10hr    2 embryo 10-12hr    1 embryo 12-14hr    2 embryo 14-16hr    2 embryo 16-18hr    3 embryo 18-20hr    2 embryo 20-22hr    4 embryo 22-24hr    5 larva L1    2 larva L2    2 larva L3 12hr old    1 larva L3 puffstage 1-2    0 larva L3 puffstage 3-6    1 larva L3 puffstage 7-9    1 white prepupae new    1 white prepupae 12hr    2 white prepupae 24hr    2 pupae 2d postWPP    2 pupae 3d postWPP    3 pupae 4d postWPP    7 adult male 01day    1 adult male 05day    1 adult male 30day    1 adult female 01day  (37) adult female 05day  (76) adult female 30day  (69) Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    76 embryo 02-04hr    9 embryo 04-06hr    5 embryo 06-08hr    2 embryo 08-10hr    2 embryo 10-12hr    1 embryo 12-14hr    2 embryo 14-16hr    2 embryo 16-18hr    3 embryo 18-20hr    2 embryo 20-22hr    4 embryo 22-24hr    5 larva L1    2 larva L2    2 larva L3 12hr old    1 larva L3 puffstage 1-2    0 larva L3 puffstage 3-6    1 larva L3 puffstage 7-9    1 white prepupae new    1 white prepupae 12hr    2 white prepupae 24hr    2 pupae 2d postWPP    2 pupae 3d postWPP    3 pupae 4d postWPP    7 adult male 01day    1 adult male 05day    1 adult male 30day    1 adult female 01day    37 adult female 05day    76 adult female 30day    69 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    76 embryo 02-04hr    9 embryo 04-06hr    5 embryo 06-08hr    2 embryo 08-10hr    2 embryo 10-12hr    1 embryo 12-14hr    2 embryo 14-16hr    2 embryo 16-18hr    3 embryo 18-20hr    2 embryo 20-22hr    4 embryo 22-24hr    5 larva L1    2 larva L2    2 larva L3 12hr old    1 larva L3 puffstage 1-2    0 larva L3 puffstage 3-6    1 larva L3 puffstage 7-9    1 white prepupae new    1 white prepupae 12hr    2 white prepupae 24hr    2 pupae 2d postWPP    2 pupae 3d postWPP    3 pupae 4d postWPP    7 adult male 01day    1 adult male 05day    1 adult male 30day    1 adult female 01day    37 adult female 05day    76 adult female 30day    69 Expression Level Scale  Extremely high  log, scaled to maximum expression level Developmental Stage   Expression Level embryo 00-02hr    76 embryo 02-04hr    9 embryo 04-06hr    5 embryo 06-08hr    2 embryo 08-10hr    2 embryo 10-12hr    1 embryo 12-14hr    2 embryo 14-16hr    2 embryo 16-18hr    3 embryo 18-20hr    2 embryo 20-22hr    4 embryo 22-24hr    5 larva L1    2 larva L2    2 larva L3 12hr old    1 larva L3 puffstage 1-2    0 larva L3 puffstage 3-6    1 larva L3 puffstage 7-9    1 white prepupae new    1 white prepupae 12hr    2 white prepupae 24hr    2 pupae 2d postWPP    2 pupae 3d postWPP    3 pupae 4d postWPP    7 adult male 01day    1 adult male 05day    1 adult male 30day    1 adult female 01day    37 adult female 05day    76 adult female 30day    69 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (76) embryo 02-04hr    9 embryo 04-06hr    5 embryo 06-08hr    2 embryo 08-10hr    2 embryo 10-12hr    1 embryo 12-14hr    2 embryo 14-16hr    2 embryo 16-18hr    3 embryo 18-20hr    2 embryo 20-22hr    4 embryo 22-24hr    5 larva L1    2 larva L2    2 larva L3 12hr old    1 larva L3 puffstage 1-2    0 larva L3 puffstage 3-6    1 larva L3 puffstage 7-9    1 white prepupae new    1 white prepupae 12hr    2 white prepupae 24hr    2 pupae 2d postWPP    2 pupae 3d postWPP    3 pupae 4d postWPP    7 adult male 01day    1 adult male 05day    1 adult male 30day    1 adult female 01day  (37) adult female 05day  (76) adult female 30day  (69) Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    76 embryo 02-04hr    9 embryo 04-06hr    5 embryo 06-08hr    2 embryo 08-10hr    2 embryo 10-12hr    1 embryo 12-14hr    2 embryo 14-16hr    2 embryo 16-18hr    3 embryo 18-20hr    2 embryo 20-22hr    4 embryo 22-24hr    5 larva L1    2 larva L2    2 larva L3 12hr old    1 larva L3 puffstage 1-2    0 larva L3 puffstage 3-6    1 larva L3 puffstage 7-9    1 white prepupae new    1 white prepupae 12hr    2 white prepupae 24hr    2 pupae 2d postWPP    2 pupae 3d postWPP    3 pupae 4d postWPP    7 adult male 01day    1 adult male 05day    1 adult male 30day    1 adult female 01day    37 adult female 05day    76 adult female 30day    69 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    76 embryo 02-04hr    9 embryo 04-06hr    5 embryo 06-08hr    2 embryo 08-10hr    2 embryo 10-12hr    1 embryo 12-14hr    2 embryo 14-16hr    2 embryo 16-18hr    3 embryo 18-20hr    2 embryo 20-22hr    4 embryo 22-24hr    5 larva L1    2 larva L2    2 larva L3 12hr old    1 larva L3 puffstage 1-2    0 larva L3 puffstage 3-6    1 larva L3 puffstage 7-9    1 white prepupae new    1 white prepupae 12hr    2 white prepupae 24hr    2 pupae 2d postWPP    2 pupae 3d postWPP    3 pupae 4d postWPP    7 adult male 01day    1 adult male 05day    1 adult male 30day    1 adult female 01day    37 adult female 05day    76 adult female 30day    69 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+    2 Schneider line 2 Sg4    9 embryonic 1182-4H    1 embryonic GM2    0 embryonic Kc167    2 embryonic S1    41 embryonic S3    2 leg disc CME L1    6 wing disc CME-W2    139 wing disc ML-DmD8    0 wing disc ML-DmD9    2 wing disc ML-DmD16-c3    1 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    1 eye-antennal disc ML-DmD11    92 antennal disc ML-DmD20-c5    54 mixed discs ML-DmD4-c1    0 CNS ML-DmBG1-c1    51 CNS ML-DmBG2-c2    14 tumorous blood cells mbn2    1 ovary fGS/OSS    524 ovary OSC    374 ovary OSS    210 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+    2 Schneider line 2 Sg4    9 embryonic 1182-4H    1 embryonic GM2    0 embryonic Kc167    2 embryonic S1  (41) embryonic S3    2 leg disc CME L1    6 wing disc CME-W2  (139) wing disc ML-DmD8    0 wing disc ML-DmD9    2 wing disc ML-DmD16-c3    1 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    1 eye-antennal disc ML-DmD11  (92) antennal disc ML-DmD20-c5  (54) mixed discs ML-DmD4-c1    0 CNS ML-DmBG1-c1  (51) CNS ML-DmBG2-c2    14 tumorous blood cells mbn2    1 ovary fGS/OSS  (524) ovary OSC  (374) ovary OSS  (210) Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Cell Line   Expression Level Schneider line 2 S2R+    2 Schneider line 2 Sg4    9 embryonic 1182-4H    1 embryonic GM2    0 embryonic Kc167    2 embryonic S1    41 embryonic S3    2 leg disc CME L1    6 wing disc CME-W2  (139) wing disc ML-DmD8    0 wing disc ML-DmD9    2 wing disc ML-DmD16-c3    1 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    1 eye-antennal disc ML-DmD11    92 antennal disc ML-DmD20-c5    54 mixed discs ML-DmD4-c1    0 CNS ML-DmBG1-c1    51 CNS ML-DmBG2-c2    14 tumorous blood cells mbn2    1 ovary fGS/OSS  (524) ovary OSC  (374) ovary OSS  (210) 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+    2 Schneider line 2 Sg4    9 embryonic 1182-4H    1 embryonic GM2    0 embryonic Kc167    2 embryonic S1    41 embryonic S3    2 leg disc CME L1    6 wing disc CME-W2    139 wing disc ML-DmD8    0 wing disc ML-DmD9    2 wing disc ML-DmD16-c3    1 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    1 eye-antennal disc ML-DmD11    92 antennal disc ML-DmD20-c5    54 mixed discs ML-DmD4-c1    0 CNS ML-DmBG1-c1    51 CNS ML-DmBG2-c2    14 tumorous blood cells mbn2    1 ovary fGS/OSS    524 ovary OSC    374 ovary OSS    210 Expression Level Scale  Extremely high  log, scaled to maximum expression level Cell Line   Expression Level Schneider line 2 S2R+    2 Schneider line 2 Sg4    9 embryonic 1182-4H    1 embryonic GM2    0 embryonic Kc167    2 embryonic S1    41 embryonic S3    2 leg disc CME L1    6 wing disc CME-W2    139 wing disc ML-DmD8    0 wing disc ML-DmD9    2 wing disc ML-DmD16-c3    1 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    1 eye-antennal disc ML-DmD11    92 antennal disc ML-DmD20-c5    54 mixed discs ML-DmD4-c1    0 CNS ML-DmBG1-c1    51 CNS ML-DmBG2-c2    14 tumorous blood cells mbn2    1 ovary fGS/OSS    524 ovary OSC    374 ovary OSS    210 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+    2 Schneider line 2 Sg4    9 embryonic 1182-4H    1 embryonic GM2    0 embryonic Kc167    2 embryonic S1  (41) embryonic S3    2 leg disc CME L1    6 wing disc CME-W2  (139) wing disc ML-DmD8    0 wing disc ML-DmD9    2 wing disc ML-DmD16-c3    1 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    1 eye-antennal disc ML-DmD11  (92) antennal disc ML-DmD20-c5  (54) mixed discs ML-DmD4-c1    0 CNS ML-DmBG1-c1  (51) CNS ML-DmBG2-c2    14 tumorous blood cells mbn2    1 ovary fGS/OSS  (524) ovary OSC  (374) ovary OSS  (210) Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Cell Line   Expression Level Schneider line 2 S2R+    2 Schneider line 2 Sg4    9 embryonic 1182-4H    1 embryonic GM2    0 embryonic Kc167    2 embryonic S1    41 embryonic S3    2 leg disc CME L1    6 wing disc CME-W2  139 wing disc ML-DmD8    0 wing disc ML-DmD9    2 wing disc ML-DmD16-c3    1 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    1 eye-antennal disc ML-DmD11    92 antennal disc ML-DmD20-c5    54 mixed discs ML-DmD4-c1    0 CNS ML-DmBG1-c1    51 CNS ML-DmBG2-c2    14 tumorous blood cells mbn2    1 ovary fGS/OSS  (524) ovary OSC  (374) ovary OSS  (210) 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+    2 Schneider line 2 Sg4    9 embryonic 1182-4H    1 embryonic GM2    0 embryonic Kc167    2 embryonic S1    41 embryonic S3    2 leg disc CME L1    6 wing disc CME-W2    139 wing disc ML-DmD8    0 wing disc ML-DmD9    2 wing disc ML-DmD16-c3    1 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    1 eye-antennal disc ML-DmD11    92 antennal disc ML-DmD20-c5    54 mixed discs ML-DmD4-c1    0 CNS ML-DmBG1-c1    51 CNS ML-DmBG2-c2    14 tumorous blood cells mbn2    1 ovary fGS/OSS    524 ovary OSC    374 ovary OSS    210 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    17 cold shock, 4-day adult    12 heat shock, 4-day adult    38 Cadmium 50 mM 6 hrs, larvae L3    0 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    26 Cadmium 100 mM 48 hrs, 4-day adult    91 Copper 0.5 mM 12 hrs, larvae L3    0 Copper 15 mM 48 hrs, 4-day adult    48 Zinc 5 mM 12 hrs, larvae L3    0 Zinc 4.5 mM 48 hrs, 4-day adult    27 Ethanol 2.5% 3 hrs, larvae L3    85 Ethanol 5% 3 hrs, larvae L3    1 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    83 Caffeine 25 mg/ml 48 hrs, 4-day adult    27 Paraquat 5 mM 48 hrs, 4-day adult    77 Paraquat 10 mM 48 hrs, 4-day adult    59 Rotenone 2 μg 12 hrs, larvae L3    1 Rotenone 8 μg 12 hrs, larvae L3    0 Expression Level Scale  Very low   Low   Moderate   Moderately high   High  Linear, scaled to Moderate expression Treatment   Expression Level extended cold, 4-day adult    17 cold shock, 4-day adult    12 heat shock, 4-day adult  (38) Cadmium 50 mM 6 hrs, larvae L3    0 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult  26 Cadmium 100 mM 48 hrs, 4-day adult  (91) Copper 0.5 mM 12 hrs, larvae L3    0 Copper 15 mM 48 hrs, 4-day adult  (48) Zinc 5 mM 12 hrs, larvae L3    0 Zinc 4.5 mM 48 hrs, 4-day adult  (27) Ethanol 2.5% 3 hrs, larvae L3  (85) Ethanol 5% 3 hrs, larvae L3    1 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult  (83) Caffeine 25 mg/ml 48 hrs, 4-day adult  (27) Paraquat 5 mM 48 hrs, 4-day adult  (77) Paraquat 10 mM 48 hrs, 4-day adult  (59) Rotenone 2 μg 12 hrs, larvae L3    1 Rotenone 8 μg 12 hrs, larvae L3    0 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Treatment   Expression Level extended cold, 4-day adult    17 cold shock, 4-day adult    12 heat shock, 4-day adult    38 Cadmium 50 mM 6 hrs, larvae L3    0 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    26 Cadmium 100 mM 48 hrs, 4-day adult    91 Copper 0.5 mM 12 hrs, larvae L3    0 Copper 15 mM 48 hrs, 4-day adult    48 Zinc 5 mM 12 hrs, larvae L3    0 Zinc 4.5 mM 48 hrs, 4-day adult    27 Ethanol 2.5% 3 hrs, larvae L3    85 Ethanol 5% 3 hrs, larvae L3    1 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    83 Caffeine 25 mg/ml 48 hrs, 4-day adult    27 Paraquat 5 mM 48 hrs, 4-day adult    77 Paraquat 10 mM 48 hrs, 4-day adult    59 Rotenone 2 μg 12 hrs, larvae L3    1 Rotenone 8 μg 12 hrs, larvae L3    0 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    17 cold shock, 4-day adult    12 heat shock, 4-day adult    38 Cadmium 50 mM 6 hrs, larvae L3    0 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    26 Cadmium 100 mM 48 hrs, 4-day adult    91 Copper 0.5 mM 12 hrs, larvae L3    0 Copper 15 mM 48 hrs, 4-day adult    48 Zinc 5 mM 12 hrs, larvae L3    0 Zinc 4.5 mM 48 hrs, 4-day adult    27 Ethanol 2.5% 3 hrs, larvae L3    85 Ethanol 5% 3 hrs, larvae L3    1 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    83 Caffeine 25 mg/ml 48 hrs, 4-day adult    27 Paraquat 5 mM 48 hrs, 4-day adult    77 Paraquat 10 mM 48 hrs, 4-day adult    59 Rotenone 2 μg 12 hrs, larvae L3    1 Rotenone 8 μg 12 hrs, larvae L3    0 Expression Level Scale  Extremely high  log, scaled to maximum expression level Treatment   Expression Level extended cold, 4-day adult    17 cold shock, 4-day adult    12 heat shock, 4-day adult    38 Cadmium 50 mM 6 hrs, larvae L3    0 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    26 Cadmium 100 mM 48 hrs, 4-day adult    91 Copper 0.5 mM 12 hrs, larvae L3    0 Copper 15 mM 48 hrs, 4-day adult    48 Zinc 5 mM 12 hrs, larvae L3    0 Zinc 4.5 mM 48 hrs, 4-day adult    27 Ethanol 2.5% 3 hrs, larvae L3    85 Ethanol 5% 3 hrs, larvae L3    1 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    83 Caffeine 25 mg/ml 48 hrs, 4-day adult    27 Paraquat 5 mM 48 hrs, 4-day adult    77 Paraquat 10 mM 48 hrs, 4-day adult    59 Rotenone 2 μg 12 hrs, larvae L3    1 Rotenone 8 μg 12 hrs, larvae L3    0 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    17 cold shock, 4-day adult    12 heat shock, 4-day adult  (38) Cadmium 50 mM 6 hrs, larvae L3    0 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult  26 Cadmium 100 mM 48 hrs, 4-day adult  (91) Copper 0.5 mM 12 hrs, larvae L3    0 Copper 15 mM 48 hrs, 4-day adult  (48) Zinc 5 mM 12 hrs, larvae L3    0 Zinc 4.5 mM 48 hrs, 4-day adult  27 Ethanol 2.5% 3 hrs, larvae L3  (85) Ethanol 5% 3 hrs, larvae L3    1 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult  (83) Caffeine 25 mg/ml 48 hrs, 4-day adult  27 Paraquat 5 mM 48 hrs, 4-day adult  (77) Paraquat 10 mM 48 hrs, 4-day adult  (59) Rotenone 2 μg 12 hrs, larvae L3    1 Rotenone 8 μg 12 hrs, larvae L3    0 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Treatment   Expression Level extended cold, 4-day adult    17 cold shock, 4-day adult    12 heat shock, 4-day adult    38 Cadmium 50 mM 6 hrs, larvae L3    0 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    26 Cadmium 100 mM 48 hrs, 4-day adult    91 Copper 0.5 mM 12 hrs, larvae L3    0 Copper 15 mM 48 hrs, 4-day adult    48 Zinc 5 mM 12 hrs, larvae L3    0 Zinc 4.5 mM 48 hrs, 4-day adult    27 Ethanol 2.5% 3 hrs, larvae L3    85 Ethanol 5% 3 hrs, larvae L3    1 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    83 Caffeine 25 mg/ml 48 hrs, 4-day adult    27 Paraquat 5 mM 48 hrs, 4-day adult    77 Paraquat 10 mM 48 hrs, 4-day adult    59 Rotenone 2 μg 12 hrs, larvae L3    1 Rotenone 8 μg 12 hrs, larvae L3    0 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    17 cold shock, 4-day adult    12 heat shock, 4-day adult    38 Cadmium 50 mM 6 hrs, larvae L3    0 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    26 Cadmium 100 mM 48 hrs, 4-day adult    91 Copper 0.5 mM 12 hrs, larvae L3    0 Copper 15 mM 48 hrs, 4-day adult    48 Zinc 5 mM 12 hrs, larvae L3    0 Zinc 4.5 mM 48 hrs, 4-day adult    27 Ethanol 2.5% 3 hrs, larvae L3    85 Ethanol 5% 3 hrs, larvae L3    1 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    83 Caffeine 25 mg/ml 48 hrs, 4-day adult    27 Paraquat 5 mM 48 hrs, 4-day adult    77 Paraquat 10 mM 48 hrs, 4-day adult    59 Rotenone 2 μg 12 hrs, larvae L3    1 Rotenone 8 μg 12 hrs, larvae L3    0 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_26 mE1_20_mRNA_expression_cluster_06
External Data & Images
Linkouts	BDGP expression data • GM05853 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0004872 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG4636-RA;CG6122-RA FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for piwi Stages(s) 1-3 Stages(s) 4-6 Stages(s) 9-10 Stages(s) 13-16
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal, with Scer\GAL4pnr-MD237 piwiGD11827 lethal | non-rescuable maternal effect piwi1 Sterility Allele female sterile piwi1 piwi2 piwi06843 female sterile, with Scer\GAL4nos.PG, Scer\GAL4nos.UTR.T:Hsim\VP16, Scer\GAL4otu.T:Hsim\VP16 piwidsRNA.shRNA.Scer\UAS.P\T female sterile | recessive piwi1 piwi06843 male fertile piwi5'T:Hsap\MYC male sterile piwi1 male sterile | recessive piwi1 Other Phenotypes Allele enhancer of variegation, with P{wA}4-4 piwi2 increased cell death (with piwi1) piwi2 increased cell death (with piwi2) piwi1 mitotic cell cycle defective | germline clone piwi1 piwi2 piwi06843 suppressor of variegation piwi1 piwi2 suppressor of variegation, with P{T1(-FRT)}T190-177 piwi1 suppressor of variegation, with P{T1}T190-177 piwi1 wild-type piwi5'T:Hsap\MYC piwi+t6.8 Phenotype manifest in Allele cephalopharyngeal skeleton | embryonic stage | maternal effect piwi1 cystoblast | non-rescuable maternal effect piwi1 dorsal appendage, with Scer\GAL4nos.PG piwidsRNA.Scer\UAS piwiGD11827 egg piwi1 egg chamber piwi2 embryo | germline clone piwi06843 piwi2 embryo | germline clone | non-rescuable maternal effect piwi1 embryonic/larval cuticle | embryonic stage | germline clone | maternal effect piwi1 embryonic head | germline clone piwi06843 piwi2 embryonic head | germline clone | non-rescuable maternal effect piwi1 female germline stem cell piwi06843 piwi1 piwi12 piwi2 female germline stem cell | cell autonomous | somatic clone piwi1 piwi2 female germline stem cell | ectopic, with Scer\GAL4hs.PB piwiEP1024 female germline stem cell | non-rescuable maternal effect piwi1 female germline stem cell | supernumerary | heat sensitive, with Scer\GAL4hs.PB piwiEP1024 germarium piwi06843 piwi1 piwi2 piwiunspecified germarium, with Scer\GAL4hs.PB piwiEP1024 germarium | non-rescuable maternal effect piwi1 germline cell | female piwiunspecified germline cell | female, with Scer\GAL4hs.PB piwiEP1024 germline cyst | non-rescuable maternal effect piwi1 male germline stem cell piwi1 nurse cell spectrosome, with Scer\GAL4hs.PB piwiEP1024 ovariole piwi06843 piwi1 piwi2 ovariole (with piwi1), with Scer\GAL4[-]: piwiEP1024 ovariole (with piwiEP1024), with Scer\GAL4[-]: piwi1 ovariole | non-rescuable maternal effect piwi1 ovary (with piwi1) piwi2 ovary (with piwi1), with Scer\GAL4[-]: piwiEP1024 ovary (with piwi2) piwi1 ovary (with piwiEP1024), with Scer\GAL4[-]: piwi1 pole cell | germline clone | maternal effect piwi1 pole cell | maternal effect | supernumerary piwi5'T:Hsap\MYC spectrosome, with Scer\GAL4hs.PB piwiEP1024 spectrosome | ectopic, with Scer\GAL4hs.PB piwiEP1024 syncytial blastoderm embryo | germline clone piwi06843 piwi2 syncytial blastoderm embryo | germline clone | non-rescuable maternal effect piwi1 testis (with piwi1), with Scer\GAL4[-]: piwiEP1024 testis (with piwiEP1024), with Scer\GAL4[-]: piwi1
Classical Alleles ( 12 )
For All Classical Alleles Show
Allele of piwi Class Mutagen Stocks Known lesion piwi06843 P-element activity 2 Yes piwi1 P-element activity 1 Yes piwi2 P-element activity 1 Yes piwie00824 piggyBac transposase 1 -- piwi12 P-element activity 0 Yes piwi4 P-element activity 0 Yes piwiCPTI003588 piggyBac transposase 0 Yes piwiEP1024 wild-type allele, loss of function allele P-element activity 0 Yes piwif07053a 0 -- piwiNt 0 Yes piwiunspecified 0 -- piwiΔ37 0 --
Alleles Carried on Transgenic Constructs ( 24 )
For All Alleles Carried on Transgenic Constructs Show
Allele of piwi Class Mutagen Stocks Known lesion piwiGD11827 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes piwiGL00626 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes piwiHMS00185 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes piwiHMS00606 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes piwiJF01394 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes piwiKK105350 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes piwi+t6.8 in vitro construct - genomic fragment 0 Yes piwi3'T:Hsap\MYC in vitro construct - coding region fusion 0 Yes piwi5'T:Hsap\MYC in vitro construct - coding region fusion 0 Yes piwiADK.T:Avic\GFP in vitro construct - coding region fusion in vitro construct - amino acid replacement 0 Yes piwibcd.3'UTR.T:Hsap\MYC in vitro construct - regulatory fusion 0 Yes piwiDAK.T:Avic\GFP in vitro construct - coding region fusion in vitro construct - amino acid replacement 0 Yes piwidsRNA.cDa in vitro construct - RNAi 0 Yes piwidsRNA.Scer\UAS in vitro construct - regulatory fusion in vitro construct - RNAi 0 Yes piwidsRNA.shRNA.Scer\UAS.P\T in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes piwihs.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes piwiNIG.6122R in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes piwiRLD in vitro construct - genomic fragment 0 Yes piwiScer\UAS.P\T.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes piwiT:Avic\GFP in vitro construct - coding region fusion 0 Yes piwiT:Zzzz\TAP in vitro construct - coding region fusion 0 Yes piwitCH322-103C03 in vitro construct - genomic fragment 0 Yes piwiV30A in vitro construct - amino acid replacement 0 Yes piwiΔNLS.T:Avic\GFP in vitro construct - coding region fusion in vitro construct - deletion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2L)BSC145 (Christensen and Cook, 2006.3.8) Df(2L)BSC213 (Christensen and Cook, 2006.8.30) Df(2L)Δ37 (Zallen et al., 2002)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD11827} NA P{KK105350} NA P{NIG.6122R} NA P{TRiP.GL00626} NA P{TRiP.HMS00185} NA P{TRiP.HMS00606} NA P{TRiP.JF01394} NA P{UAS-piwi.hp} NA P{UASp-piwi.HA} NA P{UASp-piwi.shRNA} NA heat-shock construct P{hs-myc-piwi} NA characterization construct P{5'-myc-piwi} NA P{GFP-piwi.ADK} NA P{GFP-piwi.DAK} NA P{GFP-piwi.wt} NA P{GFP-piwi.ΔNLS} NA P{piwi-3'-myc} NA P{piwi.RLD} NA P{piwi+.Rc12} NA P{piwiV30A} NA PBac{CH322-103C03} NA
Insertions
	Type of insertions Name Expression data insertion of mobile activating element P{EP}piwiEP1024 NA miscellaneous insertions P{ry11}piwi2 NA PBac{768.FSVS-0}piwiCPTI003588 NA PBac{RB}piwie00824 NA PBac{WH}piwif07053a NA insertion of enhancer trap P{lacZ}piwiNt No P{PZ}piwi1 No P{PZ}piwi4 No P{PZ}piwi06843 No
Gene Ontology: Function, Process & Cellular Component ( 25 unique terms )
Terms Based on Experimental Evidence ( 21 terms )
Molecular Function
CV term Evidence References
protein binding inferred from physical interaction with Dcr-1 (Megosh et al., 2006) inferred from physical interaction with Fmr1 (Megosh et al., 2006) inferred from physical interaction with vas (Megosh et al., 2006) RNA binding inferred from direct assay (Brennecke et al., 2007)
Biological Process
CV term Evidence References
chromatin silencing inferred from mutant phenotype (Pal-Bhadra et al., 2002, Pal-Bhadra et al., 2004, Grimaud et al., 2006) female germ-line stem cell division inferred from mutant phenotype (assigned by UniProtKB) (Cox et al., 1998) germ-line stem cell maintenance inferred from direct assay (assigned by UniProtKB) (Cox et al., 2000) heterochromatin organization involved in chromatin silencing inferred from mutant phenotype (Sienski et al., 2012) male germ-line stem cell division inferred from mutant phenotype (assigned by UniProtKB) (Cox et al., 1998) negative regulation of transposition inferred from mutant phenotype (Wang and Elgin, 2011, Klenov et al., 2011, Sienski et al., 2012) negative regulation of transposition, DNA-mediated inferred from mutant phenotype (Kalmykova et al., 2005) pole cell formation inferred from mutant phenotype (Megosh et al., 2006) positive regulation of nuclear-transcribed mRNA poly(A) tail shortening inferred from mutant phenotype (Rouget et al., 2010) posttranscriptional gene silencing inferred from mutant phenotype (Pal-Bhadra et al., 2002) stem cell division inferred from mutant phenotype (assigned by UniProtKB) (Cox et al., 1998)
Cellular Component
CV term Evidence References
chromocenter inferred from direct assay (Brower-Toland et al., 2007) cytoplasm inferred from direct assay (Zamparini et al., 2011) euchromatin inferred from direct assay (Brower-Toland et al., 2007) nucleoplasm inferred from direct assay (Lau et al., 2009) inferred from direct assay (assigned by UniProtKB) (Cox et al., 2000) nucleus inferred from direct assay (Brennecke et al., 2007, Saito et al., 2006, Grimaud et al., 2006, Zamparini et al., 2011, Olivieri et al., 2012) polytene chromosome inferred from direct assay (Brower-Toland et al., 2007) telomeric heterochromatin inferred from direct assay (Brower-Toland et al., 2007) Yb body inferred from physical interaction with armi (Handler et al., 2011)
Terms Based on Predictions or Assertions ( 8 terms )
Molecular Function
CV term Evidence References
mRNA binding non-traceable author statement (Morgan, 2002)
Biological Process
CV term Evidence References
cell fate determination traceable author statement (Deng and Lin, 2001) germ-line stem cell division traceable author statement (Deng and Lin, 2001, Lin, 2002) germ-line stem cell maintenance traceable author statement (Williams and Rubin, 2002) RNA interference traceable author statement (Cerutti, 2003)
Cellular Component
CV term Evidence References
cytoplasm non-traceable author statement (Morgan, 2002) nucleoplasm non-traceable author statement (Deng and Lin, 2001) traceable author statement (Lin, 2002) nucleus traceable author statement (Cerutti, 2003)
Sequence Ontology: Class of Gene
	nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
protein-protein
Interacting group Assay References
piwi - armi western blot, anti bait coimmunoprecipitation (Handler et al., 2011) piwi - qin anti tag coimmunoprecipitation, western blot (Anand and Kai, 2012) piwi - vret western blot, anti bait coimmunoprecipitation, anti tag coimmunoprecipitation (Handler et al., 2011, Zamparini et al., 2011)
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions piwi allele Gene References piwi2 CG7816 (Smulders-Srinivasan and Lin, 2003) hh (King et al., 2001) l(3)99Da (Smulders-Srinivasan and Lin, 2003) l(3)99De (Smulders-Srinivasan and Lin, 2003) l(3)99Dg (Smulders-Srinivasan and Lin, 2003) l(3)99Ea (Smulders-Srinivasan and Lin, 2003) RpS3 (Smulders-Srinivasan and Lin, 2003) sima (Smulders-Srinivasan and Lin, 2003) Sry-δ (Smulders-Srinivasan and Lin, 2003) tgo (Smulders-Srinivasan and Lin, 2003) piwi12 sima (Smulders-Srinivasan and Lin, 2003)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 60588 DroID - A comprehensive database of gene and protein interactions. • FBgn0004872 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 34521
Orthologs
OrthoDB Orthologs (73) - 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 EOG6CC4KV Dipteran inclusive ortholog search No orthologs identified Insect inclusive ortholog search EOG61JX76 Arthropod inclusive ortholog search EOG66DJHQ Metazoa inclusive ortholog search EOG647TM6
Orthologs in Drosophila Species (EOG6CC4KV)
Organism Common Name Gene AAA Syntenic Ortholog Multiple Dmel Genes in this Orthologous Group Drosophila melanogaster fruit fly  Dmel\piwi     Drosophila simulans   Dsim\GD22206 Y   Drosophila sechellia   Dsec\GM11236 Y   Drosophila erecta   Dere\GG10337 Y   Drosophila yakuba   Dyak\GE13110 Y   Drosophila ananassae   Dana\GF14047     Drosophila pseudoobscura pseudoobscura   Dpse\GA19370 Y   Drosophila persimilis   Dper\GL19019     Drosophila willistoni   Dwil\GK14889 Y   Drosophila virilis   Dvir\GJ13916 Y   Drosophila mojavensis   Dmoj\GI20575 Y   Drosophila grimshawi   Dgri\GH10762 Y
Orthologs in non-Drosophila Dipterans (None identified)
No non-Drosophilid orthologies identified
Orthologs in non-Dipteran Insects (EOG61JX76)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Apis mellifera Western honey bee  Amel\GB54204 Y Apis mellifera Western honey bee  Amel\GB45789 Y Nasonia vitripennis Parasitic wasp  Nvit\Nasvi2EG001968 Y Nasonia vitripennis Parasitic wasp  Nvit\Nasvi2EG004772 Y Nasonia vitripennis Parasitic wasp  Nvit\Nasvi2EG013964 Y Acromyrmex echinatior Panamanian leafcutter ant  Aech\AECH14700 Y Acromyrmex echinatior Panamanian leafcutter ant  Aech\AECH26794 Y Atta cephalotes Leafcutter ant  Acep\ACEP24111 Y Atta cephalotes Leafcutter ant  Acep\ACEP17982 Y Camponotus floridanus Florida carpenter ant  Cflo\CFLO13958 Y Camponotus floridanus Florida carpenter ant  Cflo\CFLO20073 Y Harpegnathos saltator Jerdons jumping ant  Hsal\HSAL15178 Y Harpegnathos saltator Jerdons jumping ant  Hsal\HSAL21205 Y Harpegnathos saltator Jerdons jumping ant  Hsal\HSAL15014 Y Linepithema humile Argentine ant  Lhum\LH26230 Y Linepithema humile Argentine ant  Lhum\LH19600 Y Pogonomyrmex barbatus Red harvester ant  Pbar\PB14168 Y Pogonomyrmex barbatus Red harvester ant  Pbar\PB24372 Y Solenopsis invicta Red fire ant  Sinv\SINV18536 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI006961 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI004735 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI008448 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI008672 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI008719 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI005740 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI005423 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI004068 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI30861 Y Acyrthosiphon pisum Pea aphid  Apim\ACYPI003869 Y Bombyx mori Silkmoth  Bmor\SIWI Y Pediculus humanus Human body louse  Phum\PHUM563960 Y Tribolium castaneum Red flour beetle  Tcas\piwi Y
Orthologs in non-Insect Arthropods (EOG66DJHQ)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Daphnia pulex Water flea  Dpux\nadhF Y Daphnia pulex Water flea  Dpux\JGI_V11_315989 Y Daphnia pulex Water flea  Dpux\petN Y Daphnia pulex Water flea  Dpux\JGI_V11_442507 Y Daphnia pulex Water flea  Dpux\JGI_V11_442506 Y Daphnia pulex Water flea  Dpux\JGI_V11_442508 Y Ixodes scapularis Deer tick  Isca\ISCW004464 Y
Orthologs in non-Arthropod Metazoa (EOG647TM6)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Caenorhabditis elegans Nematode  Cele\prg-1 Y Caenorhabditis elegans Nematode  Cele\prg-2 Y Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_016689gn Y Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_024002gn Y Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_023335gn Y Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_012335gn Y Danio rerio Zebrafish  Drer\piwil1 Y Danio rerio Zebrafish  Drer\ENSDARG00000062601 Y Xenopus tropicalis Western clawed frog  Xtro\piwil4 Y Xenopus tropicalis Western clawed frog  Xtro\piwil2 Y Xenopus tropicalis Western clawed frog  Xtro\piwil3 Y Gallus gallus Domestic chicken  Ggal\PIWIL1 Y Mus musculus House mouse  Mmus\Piwil1 Y Mus musculus House mouse  Mmus\Piwil4 Y Mus musculus House mouse  Mmus\Piwil2 Y Rattus norvegicus Norway rat  Rnor\Piwil1 Y Rattus norvegicus Norway rat  Rnor\ENSRNOG00000009043 Y Rattus norvegicus Norway rat  Rnor\Piwil2 Y Homo sapiens Human  Hsap\PIWIL4 Y Homo sapiens Human  Hsap\PIWIL3 Y Homo sapiens Human  Hsap\PIWIL2 Y Homo sapiens Human  Hsap\PIWIL1 Y
Human Orthologs (4)
Gene OMIM HGNC Hsap\PIWIL4     Hsap\PIWIL3     Hsap\PIWIL2     Hsap\PIWIL1
AAA Orthologs (9) based on analysis using Dmel annotation version 4.3
Organism Gene Drosophila simulans Dsim\GD22206 Drosophila sechellia Dsec\GM11236 Drosophila erecta Dere\GG10337 Drosophila yakuba Dyak\GE13110 Drosophila pseudoobscura pseudoobscura Dpse\GA19370 Drosophila willistoni Dwil\GK14889 Drosophila virilis Dvir\GJ13916 Drosophila mojavensis Dmoj\GI20575 Drosophila grimshawi Dgri\GH10762
Stocks & Reagents
Stocks Listed in FlyBase ( 11 )
Bloomington	43319 w1118; P{ry11}piwi2/CyO 43637 w1118; P{PZ}piwi1/CyO, w+* 12225 P{PZ}piwi06843 cn1/CyO; ry506 33724 y1 sc* v1; P{TRiP.HMS00606}attP2 37483 y1 sc* v1; P{TRiP.GL00626}attP40 31610 y1 v1; P{TRiP.JF01394}attP2/TM3, Ser1 34866 y1 sc* v1; P{TRiP.HMS00185}attP2/TM3, Sb1
Harvard	- PBac{RB}piwie00824
Kyoto	108882 P{PZ}piwi06843 cn1/CyO; ry506
VDRC	v101658 P{KK105350}VIE-260B v22235 w1118; P{GD11827}v22235
Genomic Clones ( 2 )
	BACR03D06 BACR20O15 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 82 )
	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	GM05853 RE21038
Other clones	FI04411
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	GM28157 LD03506 LD33002 LD35072 RE03347 RE09253 RE38167 RE59653
Other clones	66976 CK01.20A.E2 CK01.65D.H11 CK01.77B.B10 Dmel_dig_020124_0682_2763 EC255169 EK018737 EK053630 EK060437 EK194863 FGC013E07 FGC016F11 FGC018H08 FGC023D06 FGM223G03 LD03429 RP003002961 RP003031502 RP003044337 RP003063400 RP003065591 RP003066862 RP003100732 RP003107169 RP003112286 RP003112809 RP003138007 RP003144984 RP003148061 RP003148850 RP003155672 RP003169527 RP003174414 RP003197232 RP003226106 RP003244531 RP003245574 RP003249834 RP003251111 RP003251233 RP003251745 RP003268780 RP003293612 RP003295923 RP003305605 RP003324054 RP003326121 RP003326431 RP003342844 RP003348712 RP003350678 RP003355680 RP003373722 RP003385843 RP003427138 RP003450984 RP003458459 RP003470980 RP003472094 RP003486770 RP003497034 RP003514111 RP003546473 RP003548383 RP003560537 RP003601404 RP003623035 RP003644990 RP003649045 RP003654044 RP003662280
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0004872 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 34521
Antibody Information
	polyclonal (Klattenhoff et al., 2009)
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of	Source for identity of: piwi CG6122 (Gong, 1999.11)
Source for database merge of
Additional comments
Other Comments
	piwi-RISC (RNA-induced silencing complex) mediates silencing of transposable elements at the transcriptional level, and this is accompanied by local heterochromatin formation. (Sienski et al., 2012) piwi is required for maintenance of long-distance chromosomal interactions between endogenous PcG target loci. (Grimaud et al., 2006) piwi protein is localised to the nucleus and is specifically associated with rasiRNAs (repeat-associated small RNAs) derived from various repetitive elements in the genome such as retrotransposons and heterochromatic regions in vivo and shows shows target RNA cleavage ("Slicer") activity in the presence of a single-stranded guide siRNA in vitro. (Saito et al., 2006) Expression is enriched in embryonic gonads. (Shigenobu et al., 2006) piwi is required for the accumulation of repeat-associated small interfering RNAs (rasiRNAs). (Vagin et al., 2006) The piwi product binds directly to repeat-associated small interfering RNAs (rasiRNAs). (Vagin et al., 2006) piwi is required for the silencing of retrotransposons in the male germline. (Kalmykova et al., 2005) bam is epistatic to somatic niche piwi function in the ovary, but germline piwi is epistatic to bam function in the ovary. (Szakmary et al., 2005) Mutations in piwi or aub result in partial loss of the methyl-K9 form of His3, most evident at minor sites within the euchromatic arms. (Pal-Bhadra et al., 2004) piwi, aub and spn-E are suppressors of repeat-induced silencing, as seen at heterochromatic tandem arrays of insertions of the P{lacW} transposon. (Pal-Bhadra et al., 2004) piwi has a cell autonomous function in promoting stem cell division in the germline. (Cox et al., 2000) piwi mediates a somatic signalling mechanism required for the asymmetric division of germ-line stem cells to produce and maintain a daughter germ-line stem cell but is not essential for the further differentiation of the committed daughter cell. (Cox et al., 1998) Mutant analysis demonstrates the piwi function is required both to maintain germline stem cells and subsequently for the division and differentiation of the stem cell progeny in both sexes. Mutations abolish the proliferation ability of both female and male germline stem cells. (Lin and Spradling, 1997) piwi gene product affects the asymmetric division of the male and female germline stem cell (GSC). (Lin and Spradling, 1992)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_057527 RefSeq (Proteins) • NP_476875 DDBJ / EMBL / GenBank • AA802575 AAD08704 AAD08705 AAF53043 AC092226 AF104354 AF104355 AQ025014 AW941950 BI214891 CZ478664 Entrez Gene - A searchable database of RefSeq genes. • 34521 UniProtKB/Swiss-Prot • Q9VKM1
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Argonaute/Dicer protein, PAZ (IPR003100) Stem cell self-renewal protein Piwi (IPR003165) Ribonuclease H-like domain (IPR012337) BDGP expression data - Patterns of gene expression in Drosophila embryogenesis • GM05853
Linkouts
	BioGRID - A database of protein and genetic interactions • 60588 DroID - A comprehensive database of gene and protein interactions. • FBgn0004872 DRSC - Results from RNAi screens. • FBgn0004872 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0004872 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG4636-RA;CG6122-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0004872 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 34521 Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /hjmuller/piwi1.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 34521 modMine - Data generated by the modENCODE project. • FBgn0004872
Synonyms & Secondary IDs ( 5 )
Reported As
Symbol Synonym	CG6122 (Keleman et al., 2009.8.5, Ni et al., 2008, Ni et al., 2010.12.1, Ni et al., 2011.7.19) Piwi (Czech et al., 2008, Zhang et al., 2011, Zamore, 2007, Antoniewski et al., 2007, Lei, 2007, Nystul and Spradling, 2006, Dorner et al., 2006, Pane et al., 2011, Klenov et al., 2007, Prasanth and Spector, 2007, Gunawardane et al., 2007, Lin, 2007, Gunawardane et al., 2007, Zamore, 2006, Carthew, 2006, Lei et al., 2008, Shpiz et al., 2008, Yin and Lin, 2007, Saito et al., 2007, Miyoshi et al., 2009, Sato et al., 2011, Kawamura et al., 2008, Peng and Karpen, 2007, Anand and Kai, 2012, Li et al., 2009, Nishida et al., 2007, Klattenhoff et al., 2009, Gim et al., 2001, Nishida et al., 2009, Kirino et al., 2010, Khurana et al., 2011, Preall et al., 2012, Olivieri et al., 2012) PIWI (Grewal and Elgin, 2007, Lowell, 2000, Morgan, 2002, Grimaud et al., 2006, Megosh et al., 2006, Sofola et al., 2007, Dufourt et al., 2011, Bai et al., 2007, Klenov, 2008, Lin, 2008, Rehwinkel et al., 2006, Kolaczkowski et al., 2011, Mendez et al., 2011, Zhou et al., 2007) piwi (Saito et al., 2006, Ronsseray et al., 2007, Kavi and Birchler, 2007, Glaser et al., 2007, Bhadra et al., 2007, Kruger, 2007, Zaratiegui, 2007, Lei and Corces, 2006, Vagin et al., 2006, Vagin et al., 2006, Megosh et al., 2006, Shigenobu et al., 2006, Christensen and Cook, 2006.8.30, Wu and Silverman, 2007, Desset et al., 2008, Parrott et al., 2011, Lau et al., 2009, Sofola et al., 2007, Shpiz et al., 2007, Shpiz et al., 2011, Kalmykova et al., 2005, Pek and Kai, 2011, Haynes et al., 2006, Kavi et al., 2006, Brennecke et al., 2007, Payne and Braun, 2006, Kalmykova et al., 2008, Chau et al., 2008, Ronsseray et al., 2008, Kavi and Birchler, 2008, Gonzalez-Reyes, 2003, Simmons et al., 2007, Brower-Toland et al., 2007, Georlette et al., 2007, Josse et al., 2007, Xing et al., 2007, Liu et al., 2011, Tchurikov and Kretova, 2011, Qi et al., 2011, Kavi and Birchler, 2009, Belinco et al., 2009, Smulders-Srinivasan et al., 2010, Castillo et al., 2011, Yang et al., 2007, Berdnik et al., 2008, McGraw et al., 2008, Blumenstiel et al., 2008, Xie et al., 2005, Chau et al., 2009, Kirino et al., 2009, Szakmary et al., 2009, Malone et al., 2009, Shpiz et al., 2009, Habayeb et al., 2009, Chambeyron et al., 2008, Lei and Corces, 2006, Cross and Simmons, 2008, Robine et al., 2009, Venken et al., 2009, Saito et al., 2009, Lu and Clark, 2010, Fernández-Ayala et al., 2010, Moshkovich and Lei, 2010, Simmons et al., 2010, Olivieri et al., 2010, Li and Lu, 2009, Haase et al., 2010, Nagao et al., 2010, Janic et al., 2010, Jepson and Reenan, 2009, Gangaraju et al., 2011, Poulton et al., 2011, Ni et al., 2011, Pelisson et al., 2007, Saito et al., 2010, Handler et al., 2011, Liu et al., 2010, Stefanov et al., 2010, Rouget et al., 2010, Zamparini et al., 2011, Thomsen et al., 2010, Chotkowski et al., 2008, Musters et al., 2006, Wang and Elgin, 2011, Sienski et al., 2012, Klenov et al., 2011, Japanese National Institute of Genetics, 2012.5.21, Meier et al., 2012, Siddiqui et al., 2012)
Name Synonym	P-element induced wimpy testis (Lin and Spradling, 1997) piwi (Kavi et al., 2006, Josse et al., 2007, Xing et al., 2007, Cross and Simmons, 2008, Saito et al., 2009) Piwi (Carmi, 2006) PIWI (Seydoux and Braun, 2006, Ronsseray et al., 2007)
Secondary FlyBase IDs
References ( 248 )
Generate a list of	All references relating to this gene ( 248 )
List References by type	Research paper  ( 129 ) Supplementary material  ( 6 ) Review  ( 57 ) Personal communication to FlyBase  ( 12 ) Abstract  ( 34 ) FlyBase analysis  ( 3 ) Conference report  ( 4 ) Stock list  ( 1 ) Curated genome annotation  ( 1 ) Website  ( 1 )
Recent research papers ( 32 )
	Kibanov et al., 2013, Anal. Biochem. 436(1): 55--64 Multicolor fluorescence imaging of whole-mount Drosophila testes for studying spermatogenesis. [FBrf0221049] Le Thomas et al., 2013, Genes Dev. 27(4): 390--399 Piwi induces piRNA-guided transcriptional silencing and establishment of a repressive chromatin state. [FBrf0220858] Anand and Kai, 2012, EMBO J. 31(4): 870--882 The tudor domain protein Kumo is required to assemble the nuage and to generate germline piRNAs in Drosophila. [FBrf0217541] Lee and Langley, 2012, Genetics 192(4): 1411--1432 Long-term and short-term evolutionary impacts of transposable elements on Drosophila. [FBrf0220133] Meier et al., 2012, PLoS Genet. 8(5): e1002676 LINT, a Novel dL(3)mbt-Containing Complex, Represses Malignant Brain Tumour Signature Genes. [FBrf0218194] Menon and Meller, 2012, Genetics 191(3): 1023--1028 A Role for siRNA in X-Chromosome Dosage Compensation in Drosophila melanogaster. [FBrf0218921] Olivieri et al., 2012, Mol. Cell 47(6): 954--969 The Cochaperone Shutdown Defines a Group of Biogenesis Factors Essential for All piRNA Populations in Drosophila. [FBrf0219572] Preall et al., 2012, RNA 18(8): 1446--1457 shutdown is a component of the Drosophila piRNA biogenesis machinery. [FBrf0218942] Siddiqui et al., 2012, Genome Biol. 13(2): R11 Genome-wide analysis of the maternal-to-zygotic transition in Drosophila primordial germ cells. [FBrf0218847] Sienski et al., 2012, Cell 151(5): 964--980 Transcriptional silencing of transposons by piwi and maelstrom and its impact on chromatin state and gene expression. [FBrf0220033] Castillo et al., 2011, BMC Evol. Biol. 11: 258 Molecular evolution under increasing transposable element burden in Drosophila: A speed limit on the evolutionary arms race. [FBrf0216257] Dufourt et al., 2011, DNA Res. 18(6): 451--461 Polycomb group-dependent, heterochromatin protein 1-independent, chromatin structures silence retrotransposons in somatic tissues outside ovaries. [FBrf0216699] Gangaraju et al., 2011, Nat. Genet. 43(2): 153--158 Drosophila Piwi functions in Hsp90-mediated suppression of phenotypic variation. [FBrf0212873] Handler et al., 2011, EMBO J. 30(19): 3977--3993 A systematic analysis of Drosophila TUDOR domain-containing proteins identifies Vreteno and the Tdrd12 family as essential primary piRNA pathway factors. [FBrf0216344] Khurana et al., 2011, Cell 147(7): 1551--1563 Adaptation to P Element Transposon Invasion in Drosophila melanogaster. [FBrf0217035] Klenov et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(46): 18760--18765 Separation of stem cell maintenance and transposon silencing functions of Piwi protein. [FBrf0216776] Kolaczkowski et al., 2011, Mol. Biol. Evol. 28(2): 1033--1042 Recurrent adaptation in RNA interference genes across the Drosophila phylogeny. [FBrf0212798] Liu et al., 2011, Development 138(9): 1863--1873 PAPI, a novel TUDOR-domain protein, complexes with AGO3, ME31B and TRAL in the nuage to silence transposition. [FBrf0213491] Mendez et al., 2011, Chembiochem 12(7): 1084--1096 The HP1a Disordered C Terminus and Chromo Shadow Domain Cooperate to Select Target Peptide Partners. [FBrf0213514] Moshkovich et al., 2011, Genes Dev. 25(16): 1686--1701 RNAi-independent role for Argonaute2 in CTCF/CP190 chromatin insulator function. [FBrf0214686] Ni et al., 2011, Nat. Methods 8(5): 405--407 A genome-scale shRNA resource for transgenic RNAi in Drosophila. [FBrf0213581] Pane et al., 2011, EMBO J. 30(22): 4601--4615 The Cutoff protein regulates piRNA cluster expression and piRNA production in the Drosophila germline. [FBrf0217070] Parrott et al., 2011, PLoS ONE 6(9): e25087 Nucleoporin98-96 Function Is Required for Transit Amplification Divisions in the Germ Line of Drosophila melanogaster. [FBrf0216255] Pek and Kai, 2011, Proc. Natl. Acad. Sci. U.S.A. 108(29): 12007--12012 DEAD-box RNA helicase Belle/DDX3 and the RNA interference pathway promote mitotic chromosome segregation. [FBrf0214405] Poulton et al., 2011, Development 138(9): 1737--1745 The microRNA pathway regulates the temporal pattern of Notch signaling in Drosophila follicle cells. [FBrf0213494] Qi et al., 2011, J. Biol. Chem. 286(5): 3789--3797 The Yb Body, a Major Site for Piwi-associated RNA Biogenesis and a Gateway for Piwi Expression and Transport to the Nucleus in Somatic Cells. [FBrf0212916] Sato et al., 2011, Genes Dev. 25(22): 2361--2373 Maelstrom coordinates microtubule organization during Drosophila oogenesis through interaction with components of the MTOC. [FBrf0216748] Shpiz et al., 2011, Nucleic Acids Res. 39(20): 8703--8711 Mechanism of the piRNA-mediated silencing of Drosophila telomeric retrotransposons. [FBrf0216525] Tchurikov and Kretova, 2011, PLoS ONE 6(7): e21882 Both piRNA and siRNA Pathways Are Silencing Transcripts of the Suffix Element in the Drosophila melanogaster Germline and Somatic Cells. [FBrf0214474] Wang and Elgin, 2011, Proc. Natl. Acad. Sci. U.S.A. 108(52): 21164--21169 Drosophila Piwi functions downstream of piRNA production mediating a chromatin-based transposon silencing mechanism in female germ line. [FBrf0217081] Zamparini et al., 2011, Development 138(18): 4039--4050 Vreteno, a gonad-specific protein, is essential for germline development and primary piRNA biogenesis in Drosophila. [FBrf0214783] Zhang et al., 2011, Mol. Cell 44(4): 572--584 Heterotypic piRNA Ping-Pong Requires Qin, a Protein with Both E3 Ligase and Tudor Domains. [FBrf0216809] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2011 Show reviews ...