FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\piwi
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General Information
Symbol
Dmel\piwi
Species
D. melanogaster
Name
P-element induced wimpy testis
Annotation Symbol
CG6122
Feature Type
protein_coding_gene
FlyBase ID
FBgn0004872
Gene Model Status
Current
Stock Availability
21 publicly available
Gene Summary
Acts via the piwi-interacting RNA (piRNA) metabolic process, which mediates the repression of transposable elements during meiosis by forming complexes composed of piRNAs and Piwi proteins and governs the methylation and subsequent repression of transposons (PubMed:15817569, PubMed:17346786, PubMed:26808625). Directly binds piRNAs, a class of 24 to 30 nucleotide RNAs that are generated by a Dicer-independent mechanism and are primarily derived from transposons and other repeated sequence elements (PubMed:16882972). In ovarian somatic cells, mediates silencing of transposable elements at the transcriptional level in a mael-dependent manner (PubMed:23159368, PubMed:28472469). Involved in silencing of long terminal repeat (LTR) retrotransposons in male germline (PubMed:15817569). In testis, regulates spermatogenesis together with Tudor-SN (PubMed:26808625). In germ cells, mediates silencing at both transcriptional and post-transcriptional levels and is involved in the maintenance of populations of primary and secondary piRNAs. Piwi-mediated transcriptional silencing is accompanied by the formation of His3 trimethylated on 'Lys-10' (H3K9me3) associated euchromatin and heterochromatin (PubMed:23392610, PubMed:23434410, PubMed:24906153). In ovary, associates predominantly with antisense piRNAs that contain uridine at their 5' end. Association with sense piRNAs is also observed but to a lesser extent. Mediates a somatic signaling mechanism required for the maintenance of germline stem cells to produce and maintain a daughter germline stem cell (PubMed:10631171, PubMed:16949822, PubMed:9199372, PubMed:9851978). It is not essential for the further differentiation of the committed daughter cell (PubMed:9851978). Acts cell autonomously to promote germline stem cell division (PubMed:10631171, PubMed:9851978). Its role in stem cell maintenance does not seem to require nuclear localization. Required maternally for the posterior localization of osk and vas and for pole cell formation during oogenesis and early embryogenesis (PubMed:16949822). Together with Hop and Hsp83, mediates canalization, also known as developmental robustness, likely via epigenetic silencing of existing genetic variants and suppression of transposon-induced new genetic variation (PubMed:21186352). Shows RNA cleavage activity, although is not required for any of its known functions (PubMed:16882972, PubMed:23297219, PubMed:9199372). In the ovaries, forms a complex with nxf2, Panx and Nxt1 which acts as effectors of cotranscriptional transposon silencing (PubMed:31368590, PubMed:31384064). (UniProt, Q9VKM1)
Contribute a Gene Snapshot for this gene.
All Summaries
Key Links
Genomic Location
Cytogenetic map
32C1-32C1
Sequence location
Recombination map
2-44
RefSeq locus
NT_033779 REGION:10982205..10987420
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (34 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
enables chromatin DNA binding
inferred from direct assay
(Ilyin et al., 2017)
enables piRNA binding
inferred from direct assay
(Yamashiro et al., 2020, Munafò et al., 2019, Sienski et al., 2015, Brennecke et al., 2007, Saito et al., 2007)
inferred from mutant phenotype
(Yamashiro et al., 2020)
enables protein binding
inferred from physical interaction with UniProtKB:Q9V3H8,UniProtKB:Q9VV73,UniProtKB:Q9W2H9
(Batki et al., 2019, Murano et al., 2019)
inferred from physical interaction with UniProtKB:A0A0B4K7J2
(Parikh et al., 2018)
inferred from physical interaction with UniProtKB:Q9VQ91
(Zhang et al., 2018)
inferred from physical interaction with UniProtKB:Q9U3V9
(Ilyin et al., 2017)
inferred from physical interaction with UniProtKB:Q9VWE6
(Ilyin et al., 2017)
inferred from physical interaction with UniProtKB:Q9W1F4
(Ilyin et al., 2017)
inferred from physical interaction with FLYBASE:Fmr1; FB:FBgn0028734
(Megosh et al., 2006)
inferred from physical interaction with FLYBASE:Dcr-1; FB:FBgn0039016
(Megosh et al., 2006)
inferred from physical interaction with FLYBASE:vas; FB:FBgn0283442
(Megosh et al., 2006)
enables RNA binding
inferred from direct assay
(Ilyin et al., 2017)
enables RNA endonuclease activity
inferred from direct assay
(Saito et al., 2006)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
enables nucleic acid binding
inferred from electronic annotation with InterPro:IPR003165, InterPro:IPR036397
(InterPro Project Members, 2004-)
enables piRNA binding
inferred from biological aspect of ancestor with PANTHER:PTN007716312
(GO Reference Genome Project, 2011-)
enables RNA binding
inferred from electronic annotation with InterPro:IPR003100
(InterPro Project Members, 2004-)
enables RNA endonuclease activity
inferred from sequence or structural similarity with UniProtKB:A8D8P8
(Gene Ontology Curators, 2002-)
inferred from biological aspect of ancestor with PANTHER:PTN008584027
(GO Reference Genome Project, 2011-)
Biological Process (16 terms)
Terms Based on Experimental Evidence (15 terms)
CV Term
Evidence
References
involved_in female germ-line stem cell asymmetric division
inferred from mutant phenotype
(Cox et al., 1998)
involved_in germ-line stem cell population maintenance
inferred from direct assay
(Cox et al., 2000)
involved_in germarium-derived oocyte fate determination
inferred from mutant phenotype
(Ma et al., 2014)
involved_in heterochromatin formation
inferred from mutant phenotype
(Brower-Toland et al., 2007, Grimaud et al., 2006, Pal-Bhadra et al., 2004, Pal-Bhadra et al., 2002)
involved_in male germ-line stem cell asymmetric division
inferred from mutant phenotype
(Cox et al., 1998)
involved_in oogenesis
inferred from mutant phenotype
(Ku et al., 2016)
involved_in piRNA-mediated gene silencing by mRNA destabilization
inferred from mutant phenotype
(Rouget et al., 2010)
involved_in pole cell formation
inferred from mutant phenotype
(Megosh et al., 2006)
involved_in post-transcriptional gene silencing
inferred from mutant phenotype
(Pal-Bhadra et al., 2002)
involved_in primary piRNA processing
inferred from mutant phenotype
(Yamashiro et al., 2020, Sienski et al., 2015, Olivieri et al., 2012)
involved_in regulatory ncRNA-mediated gene silencing
inferred from mutant phenotype
(Le Thomas et al., 2013)
involved_in spermatogenesis
inferred from mutant phenotype
(Ku et al., 2016)
involved_in transposable element silencing by heterochromatin formation
inferred from mutant phenotype
(Sienski et al., 2015)
involved_in transposable element silencing by piRNA-mediated heterochromatin formation
inferred from mutant phenotype
(Bence et al., 2024, Andreev et al., 2022, Eastwood et al., 2021, Fabry et al., 2021, Iwasaki et al., 2021, Mugat et al., 2020, Chang et al., 2019, Akkouche et al., 2017, Sienski et al., 2015, Yu et al., 2015, Sienski et al., 2012, Klenov et al., 2011, Wang and Elgin, 2011)
involved_in transposable element silencing
inferred from mutant phenotype
(Ma et al., 2014, Kalmykova et al., 2005)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
involved_in piRNA processing
inferred from biological aspect of ancestor with PANTHER:PTN002339693
(GO Reference Genome Project, 2011-)
involved_in regulatory ncRNA-mediated gene silencing
inferred from biological aspect of ancestor with PANTHER:PTN001875625
(GO Reference Genome Project, 2011-)
involved_in spermatogenesis
inferred from biological aspect of ancestor with PANTHER:PTN002339693
(GO Reference Genome Project, 2011-)
Cellular Component (12 terms)
Terms Based on Experimental Evidence (12 terms)
CV Term
Evidence
References
located_in chromatin
inferred from direct assay
(Le Thomas et al., 2013)
located_in chromocenter
inferred from direct assay
(Brower-Toland et al., 2007)
located_in cytoplasm
inferred from direct assay
(Parikh et al., 2018, Ku et al., 2016)
inferred from direct assay
(Kistler et al., 2018, Sienski et al., 2015, Zamparini et al., 2011)
is_active_in cytosol
inferred from direct assay
(Munafò et al., 2019)
located_in cytosol
inferred from high throughput direct assay
(Lye et al., 2014)
inferred from direct assay
(Tindell et al., 2020, Yamashiro et al., 2020)
located_in euchromatin
inferred from direct assay
(Brower-Toland et al., 2007)
located_in germ cell nucleus
inferred from direct assay
(Kistler et al., 2018)
located_in heterochromatin
inferred from direct assay
(Brower-Toland et al., 2007)
located_in nucleoplasm
inferred from direct assay
(Lau et al., 2009)
inferred from direct assay
(Cox et al., 2000)
is_active_in nucleus
inferred from direct assay
(Vrettos et al., 2021, Munafò et al., 2019, Sienski et al., 2015, Le Thomas et al., 2013, Olivieri et al., 2012, Klenov et al., 2011, Zamparini et al., 2011, Brennecke et al., 2007, Grimaud et al., 2006, Saito et al., 2006)
located_in nucleus
inferred from direct assay
(Batki et al., 2019, Parikh et al., 2018, Ilyin et al., 2017, Ku et al., 2016)
is_active_in P granule
inferred from direct assay
(Tindell et al., 2020)
part_of protein-containing complex
inferred from direct assay
(Zhang et al., 2018)
is_active_in Yb body
inferred from direct assay
(Munafò et al., 2019)
inferred from physical interaction with FLYBASE:armi; FB:FBgn0041164
(Handler et al., 2011)
inferred from mutant phenotype
(Yamashiro et al., 2020)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN001875625
(GO Reference Genome Project, 2011-)
is_active_in P granule
inferred from biological aspect of ancestor with PANTHER:PTN007716312
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the argonaute family. Piwi subfamily. (Q9VKM1)
Protein Signatures (InterPro)
Summaries
Gene Group (FlyBase)
ARGONAUTE ENDORIBONUCLEASES -
Argonaute (AGO) proteins play an important role in small-RNA-mediated (miRNA, siRNA and piRNA) gene-silencing events and form the core constituent of the RNA-induced silencing complex (RISC). The PIWI domain of AGO proteins resembles RNase-H and accounts for their endonuclease activity that cleaves target mRNA complementary to the bound small RNA. (Adapted from PMID: 29447113).
Protein Function (UniProtKB)
Acts via the piwi-interacting RNA (piRNA) metabolic process, which mediates the repression of transposable elements during meiosis by forming complexes composed of piRNAs and Piwi proteins and governs the methylation and subsequent repression of transposons (PubMed:15817569, PubMed:17346786, PubMed:26808625). Directly binds piRNAs, a class of 24 to 30 nucleotide RNAs that are generated by a Dicer-independent mechanism and are primarily derived from transposons and other repeated sequence elements (PubMed:16882972). In ovarian somatic cells, mediates silencing of transposable elements at the transcriptional level in a mael-dependent manner (PubMed:23159368, PubMed:28472469). Involved in silencing of long terminal repeat (LTR) retrotransposons in male germline (PubMed:15817569). In testis, regulates spermatogenesis together with Tudor-SN (PubMed:26808625). In germ cells, mediates silencing at both transcriptional and post-transcriptional levels and is involved in the maintenance of populations of primary and secondary piRNAs. Piwi-mediated transcriptional silencing is accompanied by the formation of His3 trimethylated on 'Lys-10' (H3K9me3) associated euchromatin and heterochromatin (PubMed:23392610, PubMed:23434410, PubMed:24906153). In ovary, associates predominantly with antisense piRNAs that contain uridine at their 5' end. Association with sense piRNAs is also observed but to a lesser extent. Mediates a somatic signaling mechanism required for the maintenance of germline stem cells to produce and maintain a daughter germline stem cell (PubMed:10631171, PubMed:16949822, PubMed:9199372, PubMed:9851978). It is not essential for the further differentiation of the committed daughter cell (PubMed:9851978). Acts cell autonomously to promote germline stem cell division (PubMed:10631171, PubMed:9851978). Its role in stem cell maintenance does not seem to require nuclear localization. Required maternally for the posterior localization of osk and vas and for pole cell formation during oogenesis and early embryogenesis (PubMed:16949822). Together with Hop and Hsp83, mediates canalization, also known as developmental robustness, likely via epigenetic silencing of existing genetic variants and suppression of transposon-induced new genetic variation (PubMed:21186352). Shows RNA cleavage activity, although is not required for any of its known functions (PubMed:16882972, PubMed:23297219, PubMed:9199372). In the ovaries, forms a complex with nxf2, Panx and Nxt1 which acts as effectors of cotranscriptional transposon silencing (PubMed:31368590, PubMed:31384064).
(UniProt, Q9VKM1)
Summary (Interactive Fly)

a novel protein involved in stem cell renewal and asymmetric cell division - piRNA-guided slicing of transposon transcripts enforces their transcriptional silencing via specifying the nuclear piRNA repertoire - maintains germline stem cells and oogenesis in Drosophila through negative regulation of Polycomb group proteins Piwi modulates chromatin accessibility by regulating multiple factors including Histone H1 to repress transposons

Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\piwi for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q9VKM1)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
PDB - An information portal to biological macromolecular structures
Comments on Gene Model

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.50

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
piwi-RA
FBtr0080166
NM_057527
3073
843
piwi-RB
FBtr0340227
NM_001298896
2969
843
Additional Transcript Data and 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
UniProt
RefSeq ID
GenBank
piwi-PA
FBpp0079755
97.2
843
10.00
Q9VKM1
NP_476875
AAF53043
piwi-PB
FBpp0309202
97.2
843
10.00
Q9VKM1
NP_001285825
AHN54339
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

843 aa isoforms: piwi-PA, piwi-PB
Additional Polypeptide Data and Comments
Reported size (kDa)

843 (aa); 97.2 (kD predicted)

(Cox et al., 1998)
Comments
External Data
Subunit Structure (UniProtKB)

In the ovaries, part of a complex composed of at least Panx, nxf2, piwi and Nxt1 (PubMed:26472911, PubMed:26494711, PubMed:31368590, PubMed:31384064). The complex is knowns as Panx-induced co-transcriptional silencing (PICTS) complex, Panx-nxf2-dependent TAP/p15 silencing (Pandas complex), SFiNX (silencing factor interacting nuclear export variant) or piwi-Panx-nxf2-p15 (PPNP) complex (PubMed:26472911, PubMed:26494711, PubMed:31368590, PubMed:31384064). Interacts with vas; this interaction is RNA-independent (PubMed:16949822). Interacts with Dcr-1 and Fmr1; these interactions occur in polar granules (PubMed:16949822). Interacts (via N-terminal region) with CBX5 (via chromoshadow domain) (PubMed:17875665). Forms a complex with Hsp83 and Hop; probably Hop mediates the interaction between piwi and Hsp83 (PubMed:21186352). Forms a complex with Yb body components armi and fs(1)Yb; this interaction is required for proper piRNA loading and nuclear localization of piwi (PubMed:20966047). Interaction of Piwi and fs(1)Yb is likely to occur via armi (PubMed:20966047). Interacts (via the N-terminal region when unmethylated or symmetrically methylated at Arg-10) with papi (via Tudor domain) (PubMed:21447556, PubMed:29531043). Interacts with vret (PubMed:21831924). Interacts with Panx (PubMed:26472911, PubMed:26494711). Interacts with arx (PubMed:23913921, PubMed:23913922). Interacts with Tudor-SN (PubMed:26808625). Interacts with Nup358 (via N-terminus) (PubMed:29735528). Associates with the nuclear pore complex via interaction with Elys (PubMed:28472469). Interacts with thoc5; the interaction might be partly RNA-mediated (PubMed:28472469). Interacts with xmas-2 (PubMed:28472469).

(UniProt, Q9VKM1)
Post Translational Modification

Symmetrically dimethylated, most likely by csul (PubMed:19377467, PubMed:26212455, PubMed:29531043). Methylation at Arg-10 enhances binding to papi whereas methylation at Arg-7, Arg-9 or Arg-11 reduces binding affinity to papi (PubMed:29531043).

Phosphorylated on serine and tyrosine residues in an Hsp83-dependent manner.

(UniProt, Q9VKM1)
Crossreferences
InterPro - A database of protein families, domains and functional sites
PDB - An information portal to biological macromolecular structures
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\piwi using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

-0.28

Transcript Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult fat body
• nucleus
(Jones et al., 2016)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
gonad
(Parrott et al., 2011)
pre-blastoderm stage
organism
(Cox et al., 1998)
embryonic stage 1 -- 3
pole plasm

Comment: maternally deposited

(Fisher et al., 2012)
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
organism

Comment: rapidly degraded

(Fisher et al., 2012)
organism | faint | ubiquitous
(Fisher et al., 2012)
primordial germ cell
(Fisher et al., 2012)
embryonic stage 7 -- 8
primordial germ 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)
ovarian sheath
(Cox et al., 1998)
ovary
(Cox et al., 1998)
germarium cap cell
(Cox et al., 1998)
RT-PCR
Stage
Tissue/Position (including subcellular localization)
Reference
oogenesis
ovary
(Lau et al., 2009)
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
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage | male & spermatogenesis
hub cell
(Nishida et al., 2007)
spermatogenesis & adult stage | male
somatic cell of testis
(Nishida et al., 2007)
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in chromatin
inferred from direct assay
(Le Thomas et al., 2013)
located_in chromocenter
inferred from direct assay
(Brower-Toland et al., 2007)
located_in cytoplasm
inferred from direct assay
(Parikh et al., 2018, Ku et al., 2016)
inferred from direct assay
(Kistler et al., 2018, Sienski et al., 2015, Zamparini et al., 2011)
is_active_in cytosol
inferred from direct assay
(Munafò et al., 2019)
located_in cytosol
inferred from high throughput direct assay
(Lye et al., 2014)
inferred from direct assay
(Tindell et al., 2020, Yamashiro et al., 2020)
located_in euchromatin
inferred from direct assay
(Brower-Toland et al., 2007)
located_in germ cell nucleus
inferred from direct assay
(Kistler et al., 2018)
located_in heterochromatin
inferred from direct assay
(Brower-Toland et al., 2007)
located_in nucleoplasm
inferred from direct assay
(Lau et al., 2009)
inferred from direct assay
(Cox et al., 2000)
is_active_in nucleus
inferred from direct assay
(Vrettos et al., 2021, Munafò et al., 2019, Sienski et al., 2015, Le Thomas et al., 2013, Olivieri et al., 2012, Klenov et al., 2011, Zamparini et al., 2011, Brennecke et al., 2007, Grimaud et al., 2006, Saito et al., 2006)
located_in nucleus
inferred from direct assay
(Batki et al., 2019, Parikh et al., 2018, Ilyin et al., 2017, Ku et al., 2016)
is_active_in P granule
inferred from direct assay
(Tindell et al., 2020)
part_of protein-containing complex
inferred from direct assay
(Zhang et al., 2018)
is_active_in Yb body
inferred from direct assay
(Munafò et al., 2019)
inferred from physical interaction with FLYBASE:armi; FB:FBgn0041164
(Handler et al., 2011)
inferred from mutant phenotype
(Yamashiro et al., 2020)
Expression Deduced from Reporters
Reporter: PBac{768.FSVS-0}piwiCPTI003588
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult brain

Comment: punctate expression throughout adult brain

(Knowles-Barley, 2011.8.24)
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\piwi in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 4-6
  • Stages(s) 9-10
  • Stages(s) 13-16
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 24 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 77 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of piwi
Transgenic constructs containing regulatory region of piwi
Aberrations (Deficiencies and Duplications) ( 3 )
Inferred from experimentation ( 3 )
Gene 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)
Inferred from location ( 4 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal, with Scer\GAL4pnr-MD237
lethal | non-rescuable maternal effect
lethal - all die during P-stage, with Scer\GAL4Mef2.PR
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4tin.CΔ4
viable (with piwi1)
viable (with piwiDUO.Tag:MYC)
viable (with piwig1)
viable (with piwig2)
Sterility
Allele
female fertile (with Df(2R)ED3181)
female sterile
female sterile, with Scer\GAL4nanos.PG
female sterile, with Scer\GAL4VP16.nanos.UTR
female sterile, with Scer\GAL4VP16.nanos.UTR, Scer\GAL4VP16.otu, Scer\GAL4nanos.PG
female sterile (with piwi2)
female sterile (with piwi06843)
female sterile (with piwig1)
female sterile (with piwig2)
female sterile | dominant
female sterile | recessive
fertile, with piwi2/piwi1
fertile (with piwi1)
fertile (with piwi1), with piwinls.Tag:FLAG
fertile (with piwi2), with piwinls.Tag:FLAG
fertile (with piwiDUO.Tag:MYC)
male fertile
male sterile
male sterile | recessive
sterile, with piwi2/piwi1
sterile (with piwi1), with piwiYK.nls.Tag:FLAG
sterile (with piwi2), with piwiYK.nls.Tag:FLAG
Other Phenotypes
Allele
abnormal cell number | female, with Scer\GAL4NP1624
abnormal cell number | oogenesis, with Scer\GAL4bab1-Gal4
abnormal cell number | oogenesis, with Scer\GAL4GMR42G01
abnormal cell number | oogenesis, with Scer\GAL4Unc-115a-KH2261
abnormal cell number | oogenesis (with piwi2)
abnormal cell number | oogenesis (with piwi06843)
abnormal cell shape | oogenesis, with Scer\GAL4C587
abnormal mitotic cell cycle | germline clone
abnormal size | oogenesis
abnormal size | oogenesis (with piwi2)
abnormal size | oogenesis (with piwi06843)
abnormal size | oogenesis | recessive
decreased cell number | cell autonomous | germline clone | oogenesis, with Scer\GAL4VP16.nanos.-FRT
decreased cell number | female | third instar larval stage, with Scer\GAL4VP16.nanos.UTR
decreased cell number | oogenesis, with Scer\GAL4C587
decreased cell number | oogenesis, with Scer\GAL4VP16.nanos.UTR
decreased fecundity | female (with piwi2)
decreased fecundity | female (with piwi06843)
decreased fecundity | female (with piwig1)
decreased fecundity | female (with piwig2)
decreased fecundity | female | recessive
enhancer of variegation, with P{wA}4-4
increased cell death (with piwi1)
increased cell death (with piwi2)
increased cell death | oogenesis, with Scer\GAL4C587
increased cell number | cell non-autonomous | germline clone | oogenesis, with Scer\GAL4VP16.nanos.-FRT
increased cell number | oogenesis, with Scer\GAL4C587
increased cell number | oogenesis, with Scer\GAL4hs.PB
long lived | dominant
neoplasia | oogenesis (with piwi2)
neoplasia | oogenesis (with piwi06843)
non-suppressor of variegation | dominant, with P{hsp26-pt-T}8-M76
non-suppressor of variegation | dominant, with P{hsp26-pt-T}39C-66
non-suppressor of variegation | dominant, with P{w+(-1360)}1250
non-suppressor of variegation | dominant, with P{w+(FRT.attP.1360)}1250
suppressor of variegation
suppressor of variegation, with P{T1(-FRT)}T190-177
suppressor of variegation, with P{T1}T190-177
suppressor of variegation | dominant, with P{w+(FRT.attP.1360)}1250
wild-type
Phenotype manifest in
Allele
cephalopharyngeal skeleton | embryonic stage | maternal effect
cystoblast (with piwi2)
cystoblast (with piwi06843)
cystoblast | non-rescuable maternal effect
dorsal appendage, with Scer\GAL4nanos.PG
egg
egg chamber
egg chamber, with Scer\GAL4NP1624
egg chamber (with Df(2L)BSC145)
embryo | embryonic stage 4 | germline clone
embryo | embryonic stage 4 | germline clone | non-rescuable maternal effect
embryo | germline clone
embryo | germline clone | non-rescuable maternal effect
embryonic/larval cuticle | embryonic stage | germline clone | maternal effect
embryonic head | germline clone
embryonic head | germline clone | non-rescuable maternal effect
escort cell, with Scer\GAL4C587
female germline stem cell
female germline stem cell, with Scer\GAL4bab1-Gal4
female germline stem cell, with Scer\GAL4C587
female germline stem cell, with Scer\GAL4Unc-115a-KH2261
female germline stem cell (with piwi2)
female germline stem cell (with piwi06843)
female germline stem cell | cell autonomous | germline clone | oogenesis, with Scer\GAL4VP16.nanos.-FRT
female germline stem cell | cell autonomous | somatic clone
female germline stem cell | cell non-autonomous | germline clone | oogenesis, with Scer\GAL4VP16.nanos.-FRT
female germline stem cell | ectopic, with Scer\GAL4hs.PB
female germline stem cell | increased number | heat sensitive, with Scer\GAL4hs.PB
female germline stem cell | non-rescuable maternal effect
female germline stem cell | oogenesis, with Scer\GAL4VP16.nanos.UTR
germarium
germarium, with Scer\GAL4bab1-Gal4
germarium, with Scer\GAL4C587
germarium, with Scer\GAL4hs.PB
germarium, with Scer\GAL4Unc-115a-KH2261
germarium, with Scer\GAL4VP16.nanos.UTR
germarium | cell autonomous | germline clone, with Scer\GAL4VP16.nanos.-FRT
germarium | cell non-autonomous | germline clone, with Scer\GAL4VP16.nanos.-FRT
germarium | non-rescuable maternal effect
germarium cap cell, with Scer\GAL4bab1-Gal4
germline cell | female
germline cell | female, with Scer\GAL4hs.PB
germline cyst | non-rescuable maternal effect
male germline stem cell
nurse cell, with Scer\GAL4GMR42G01
nurse cell | increased number, with Scer\GAL4NP1624
nurse cell spectrosome, with Scer\GAL4hs.PB
oocyte, with Scer\GAL4NP1624
ovariole
ovariole, with Scer\GAL4GMR42G01
ovariole (with piwi1)
ovariole (with piwiEP1024)
ovariole | non-rescuable maternal effect
ovary
ovary (with piwi1)
ovary (with piwi2)
ovary (with piwi06843)
ovary (with piwiEP1024)
primordial germ cell | germline clone | maternal effect
primordial germ cell | maternal effect | increased number
spectrosome, with Scer\GAL4hs.PB
spectrosome | ectopic, with Scer\GAL4hs.PB
testis (with piwi1)
testis (with piwiEP1024)
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (8)
Hsap\PIWIL3
11 of 14
Yes
No
Hsap\PIWIL1
10 of 14
No
No
Hsap\PIWIL4
9 of 14
No
No
Hsap\PIWIL2
8 of 14
No
No
Hsap\AGO1
1 of 14
No
No
Hsap\AGO2
1 of 14
No
No
Hsap\AGO3
1 of 14
No
No
1  
Hsap\AGO4
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (7)
Rnor\Piwil1
10 of 14
Yes
No
Rnor\Piwil4
9 of 14
No
No
Rnor\Piwil2
8 of 14
No
No
Rnor\Ago1
1 of 14
No
No
Rnor\Ago2
1 of 14
No
No
Rnor\Ago3
1 of 14
No
No
Rnor\Ago4
1 of 14
No
No
Mus musculus (laboratory mouse) (7)
Mmus\Piwil1
9 of 14
Yes
No
Mmus\Piwil2
8 of 14
No
No
Mmus\Piwil4
8 of 14
No
No
Mmus\Ago1
1 of 14
No
No
Mmus\Ago2
1 of 14
No
No
Mmus\Ago3
1 of 14
No
No
Mmus\Ago4
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (8)
Xtro\piwil3
8 of 13
Yes
No
Xtro\piwil2
5 of 13
No
No
Xtro\piwil4
5 of 13
No
No
Xtro\piwil1
3 of 13
No
No
Xtro\ago2
1 of 13
No
No
Xtro\ago4
1 of 13
No
No
Xtro\cyp2d6
1 of 13
No
No
Xtro\inhbc.1
1 of 13
No
No
Danio rerio (Zebrafish) (7)
Drer\piwil1
11 of 14
Yes
No
Drer\piwil2
7 of 14
No
No
Drer\ago1
1 of 14
No
No
Drer\ago2
1 of 14
No
No
Drer\ago3a
1 of 14
No
No
Drer\ago3b
1 of 14
No
No
Drer\ago4
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (21)
Cele\prg-1
13 of 14
Yes
No
Cele\csr-1
2 of 14
No
Yes
Cele\ppw-1
2 of 14
No
Yes
Cele\sago-2
2 of 14
No
Yes
Cele\wago-1
2 of 14
No
No
Cele\wago-4
2 of 14
No
No
Cele\wago-5
2 of 14
No
No
Cele\alg-1
1 of 14
No
No
Cele\alg-2
1 of 14
No
No
Cele\alg-3
1 of 14
No
No
Cele\alg-4
1 of 14
No
No
Cele\alg-5
1 of 14
No
No
Cele\C04F12.1
1 of 14
No
No
Cele\ergo-1
1 of 14
No
No
Cele\hrde-1
1 of 14
No
No
Cele\nrde-3
1 of 14
No
No
Cele\ppw-2
1 of 14
No
No
Cele\prg-2
1 of 14
No
No
Cele\rde-1
1 of 14
No
No
Cele\sago-1
1 of 14
No
No
Cele\wago-10
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (5)
Agam\AgaP_AGAP009509
7 of 12
Yes
No
Agam\AgaP_AGAP008862
6 of 12
No
No
Agam\AgaP_AGAP011204
6 of 12
No
No
Agam\AgaP_AGAP011537
1 of 12
No
No
Agam\AgaP_AGAP011717
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (10)
Atha\AGO3
3 of 13
Yes
Yes
Atha\AGO2
2 of 13
No
No
Atha\AGO6
2 of 13
No
No
Atha\AGO7
2 of 13
No
No
Atha\AGO1
1 of 13
No
No
Atha\AGO4
1 of 13
No
No
Atha\AGO5
1 of 13
No
No
Atha\AGO9
1 of 13
No
No
Atha\AGO10
1 of 13
No
No
Atha\AT5G21030
1 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (1)
Spom\ago1
1 of 12
Yes
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:piwi. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (4)
aub
11 of 13
AGO3
10 of 13
AGO1
1 of 13
AGO2
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Allele
Disease
Evidence
References
piwiHMS00606
model of  primary ovarian insufficiency
CEA
(Gorsi et al., 2022)
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 3 )
Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
Homo sapiens (Human)
Gene name
Score
OMIM
OMIM Phenotype
DO term
Complementation?
Transgene?
PIWIL3; piwi like RNA-mediated gene silencing 3
11 of 14
610314
      PIWIL1; piwi like RNA-mediated gene silencing 1
      10 of 14
      605571
          PIWIL4; piwi like RNA-mediated gene silencing 4
          9 of 14
          610315
              PIWIL2; piwi like RNA-mediated gene silencing 2
              8 of 14
              610312
                  Functional Complementation Data
                  Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
                  Interactions
                  Summary of Physical Interactions
                  Interaction Browsers
                  View in FlyBase Interactions Browser   View in MIST tool (external link)
                  Please see the Physical Interaction reports below for full details
                  protein-protein
                  Physical Interaction
                  Assay
                  References
                  piwi - Caf1-55
                  anti bait coimmunoprecipitation, western blot
                  (Mugat et al., 2020)
                  piwi - Cp190
                  anti bait coimmunoprecipitation, western blot
                  (Moshkovich et al., 2011)
                  piwi - Dcr-1
                  anti tag coimmunoprecipitation, western blot
                  (Megosh et al., 2006)
                  piwi - E(z)
                  anti bait coimmunoprecipitation, western blot
                  (Peng et al., 2016)
                  piwi - Elys
                  anti bait coimmunoprecipitation, western blot
                  (Ilyin et al., 2017)
                  piwi - Fmr1
                  anti tag coimmunoprecipitation, western blot, anti tag western blot, pull down, anti bait coimmunoprecipitation
                  (Jiang et al., 2016, Megosh et al., 2006)
                  piwi - Gasz
                  anti tag coimmunoprecipitation, western blot
                  (Yamashiro et al., 2020)
                  piwi - HDAC1
                  anti bait coimmunoprecipitation, western blot
                  (Mugat et al., 2020)
                  piwi - Hen1
                  pull down, western blot
                  (Ohtani et al., 2013)
                  piwi - His1
                  anti bait coimmunoprecipitation, western blot, peptide massfingerprinting, pull down, anti tag western blot
                  (Iwasaki et al., 2016)
                  piwi - Hsp83
                  anti tag coimmunoprecipitation, western blot, anti bait coimmunoprecipitation
                  (Gangaraju et al., 2011)
                  piwi - Kap-α3
                  anti bait coimmunoprecipitation, anti tag western blot
                  (Yashiro et al., 2018)
                  piwi - Klp10A
                  anti bait coimmunoprecipitation, western blot, northern blot, coimmunoprecipitation, Identification by mass spectrometry
                  (Venkei et al., 2020, Saito et al., 2009)
                  piwi - MEP-1
                  anti bait coimmunoprecipitation, western blot
                  (Mugat et al., 2020)
                  piwi - Mi-2
                  anti bait coimmunoprecipitation, western blot, Identification by mass spectrometry
                  (Mugat et al., 2020, Onishi et al., 2020)
                  piwi - Nbr
                  anti tag coimmunoprecipitation, western blot, peptide massfingerprinting, anti tag western blot
                  (Hayashi et al., 2016, Wang et al., 2016)
                  piwi - Not1
                  anti bait coimmunoprecipitation, western blot
                  (Kordyukova et al., 2020)
                  piwi - Nup358
                  anti bait coimmunoprecipitation, western blot, pull down, anti tag western blot
                  (Parikh et al., 2018)
                  piwi - Nxt1
                  anti tag coimmunoprecipitation, western blot
                  (Fabry et al., 2019)
                  piwi - Panx
                  coimmunoprecipitation, western blot, anti tag coimmunoprecipitation, anti tag western blot, anti bait coimmunoprecipitation, Identification by mass spectrometry
                  (Onishi et al., 2020, Batki et al., 2019, Fabry et al., 2019, Murano et al., 2019, Sienski et al., 2015, Yu et al., 2015)
                  piwi - Ssrp
                  anti tag coimmunoprecipitation, western blot, Identification by mass spectrometry, anti bait coimmunoprecipitation
                  (Chen et al., 2025)
                  piwi - Stip1
                  anti tag coimmunoprecipitation, western blot, ion exchange chromatography, molecular sieving, peptide massfingerprinting, anti bait coimmunoprecipitation
                  (Gangaraju et al., 2011)
                  piwi - Su(var)2-10
                  pull down, anti tag western blot, anti tag coimmunoprecipitation, bimolecular fluorescence complementation, fluorescence microscopy, western blot
                  (Bence et al., 2024, Mugat et al., 2020, Ninova et al., 2020)
                  piwi - Su(var)205
                  anti bait coimmunoprecipitation, western blot, clip, reverse transcription pcr, fluorescence polarization spectroscopy, predetermined participant, two hybrid, nuclear magnetic resonance
                  (Casale et al., 2019, Mendez et al., 2013, Mendez et al., 2011, Brower-Toland et al., 2007)
                  piwi - Su(var)3-3
                  anti bait coimmunoprecipitation, western blot
                  (Lepesant et al., 2020)
                  piwi - Su(z)12
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Peng et al., 2016)
                  piwi - Sumo
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Ninova et al., 2023)
                  piwi - Tdrd3
                  anti bait coimmunoprecipitation, western blot, Identification by mass spectrometry
                  (Lee et al., 2025)
                  piwi - Tudor-SN
                  anti bait coimmunoprecipitation, western blot, ion exchange chromatography, molecular sieving, peptide massfingerprinting
                  (Ku et al., 2016)
                  piwi - U2af38
                  anti bait coimmunoprecipitation, western blot
                  (He et al., 2022)
                  piwi - armi
                  anti bait coimmunoprecipitation, western blot, affinity chromatography technology, Identification by mass spectrometry, anti tag western blot, anti tag coimmunoprecipitation
                  (Lee et al., 2025, Yamashiro et al., 2020, Munafò et al., 2019, Ilyin et al., 2017, Pandey et al., 2017, Handler et al., 2011, Olivieri et al., 2010, Saito et al., 2010)
                  piwi - arx
                  anti bait coimmunoprecipitation, peptide massfingerprinting, western blot, anti tag coimmunoprecipitation, pull down, Identification by mass spectrometry, luminescence based mammalian interactome mapping, molecular sieving, molecular weight estimation by staining
                  (Mugat et al., 2020, Onishi et al., 2020, Yamaguchi et al., 2020, Dönertas et al., 2013, Ohtani et al., 2013)
                  piwi - brm
                  anti bait coimmunoprecipitation, western blot, Identification by mass spectrometry
                  (Onishi et al., 2020)
                  piwi - brn
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Kawaguchi et al., 2025)
                  piwi - corto
                  anti bait coimmunoprecipitation, western blot
                  (Peng et al., 2016)
                  piwi - esc
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Peng et al., 2016)
                  piwi - fs(1)Yb
                  proximity-dependent biotin identification, Identification by mass spectrometry, anti bait coimmunoprecipitation, western blot
                  (Lee et al., 2025, Munafò et al., 2021)
                  piwi - mael
                  anti bait coimmunoprecipitation, anti tag western blot, western blot
                  (Onishi et al., 2020)
                  piwi - nxf2
                  anti tag coimmunoprecipitation, western blot, Identification by mass spectrometry, anti bait coimmunoprecipitation
                  (Fabry et al., 2019, Murano et al., 2019)
                  piwi - papi
                  two hybrid, anti tag coimmunoprecipitation, anti tag western blot, x-ray crystallography, isothermal titration calorimetry, predetermined participant, pull down, western blot, Identification by mass spectrometry, anti bait coimmunoprecipitation, molecular weight estimation by staining
                  (Zhang et al., 2018, Liu et al., 2011)
                  piwi - qin
                  anti tag coimmunoprecipitation, western blot
                  (Anand and Kai, 2012)
                  piwi - shu
                  anti bait coimmunoprecipitation, western blot
                  (Olivieri et al., 2012)
                  piwi - spn-E
                  anti tag coimmunoprecipitation, cross-linking study, western blot
                  (Lin et al., 2023)
                  piwi - squ
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Haase et al., 2010)
                  piwi - tej
                  anti tag coimmunoprecipitation, cross-linking study, western blot
                  (Lin et al., 2023)
                  piwi - thoc5
                  anti bait coimmunoprecipitation, western blot
                  (Ilyin et al., 2017)
                  piwi - twin
                  anti tag coimmunoprecipitation, western blot, anti tag western blot
                  (Kordyukova et al., 2020)
                  piwi - vas
                  anti tag coimmunoprecipitation, cross-linking study, western blot, anti bait coimmunoprecipitation
                  (Lin et al., 2023, Venkei et al., 2020, Megosh et al., 2006)
                  piwi - vret
                  western blot, anti bait coimmunoprecipitation, anti tag coimmunoprecipitation
                  (Handler et al., 2011, Zamparini et al., 2011)
                  piwi - xmas
                  anti bait coimmunoprecipitation, western blot
                  (Ilyin et al., 2017)
                  piwi - zuc
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Haase et al., 2010)
                  RNA-protein
                  Physical Interaction
                  Assay
                  References
                  piwi - Klp10A
                  anti bait coimmunoprecipitation, western blot, northern blot, coimmunoprecipitation, Identification by mass spectrometry
                  (Venkei et al., 2020, Saito et al., 2009)
                  piwi - Su(var)205
                  anti bait coimmunoprecipitation, western blot, clip, reverse transcription pcr, fluorescence polarization spectroscopy, predetermined participant, two hybrid, nuclear magnetic resonance
                  (Casale et al., 2019, Mendez et al., 2013, Mendez et al., 2011, Brower-Toland et al., 2007)
                  piwi - brat
                  anti bait coimmunoprecipitation, northern blot
                  (Saito et al., 2009)
                  piwi - kay
                  anti tag coimmunoprecipitation, quantitative reverse transcription pcr
                  (Klein et al., 2016)
                  piwi - lncRNA:flam
                  anti bait coimmunoprecipitation, full identification by DNA sequencing, nucleotide sequence identification
                  (Fagegaltier et al., 2016, Saito et al., 2009)
                  piwi - tj
                  anti bait coimmunoprecipitation, nucleotide sequence identification, northern blot
                  (Saito et al., 2009)
                  Summary of Genetic Interactions
                  Interaction Browsers
                  View in FlyBase Interactions Browser   View in MIST tool (external link)
                  Please look at the allele data for full details of the genetic interactions
                  Starting gene(s)
                  Interaction type
                  Interacting gene(s)
                  Reference
                  piwi
                  suppressible
                  arm
                  (Lin et al., 2020)
                  piwi
                  enhanceable
                  CycJ|armi
                  (Atikukke et al., 2014)
                  piwi
                  suppressible
                  dpp
                  (Ma et al., 2014)
                  piwi
                  suppressible
                  hh
                  (King et al., 2001)
                  piwi
                  suppressible
                  l(3)99Da
                  (Smulders-Srinivasan and Lin, 2003)
                  piwi
                  suppressible
                  l(3)99De
                  (Smulders-Srinivasan and Lin, 2003)
                  piwi
                  suppressible
                  l(3)99Dg
                  (Smulders-Srinivasan and Lin, 2003)
                  piwi
                  suppressible
                  l(3)99Ea
                  (Smulders-Srinivasan and Lin, 2003)
                  piwi
                  suppressible
                  RpS3
                  (Smulders-Srinivasan and Lin, 2003)
                  piwi
                  suppressible
                  sgg
                  (Lin et al., 2020)
                  piwi
                  suppressible
                  sima
                  (Smulders-Srinivasan and Lin, 2003)
                  piwi
                  suppressible
                  Sry-δ
                  (Smulders-Srinivasan and Lin, 2003)
                  piwi
                  suppressible
                  Ssrp
                  (Chen et al., 2025)
                  piwi
                  suppressible
                  Su(var)2-1
                  (Yang et al., 2019)
                  piwi
                  suppressible
                  Tehao
                  (Lin et al., 2020)
                  piwi
                  suppressible
                  tgo
                  (Smulders-Srinivasan and Lin, 2003)
                  piwi
                  suppressible
                  Tl
                  (Lin et al., 2020)
                  piwi
                  suppressible
                  Tudor-SN
                  (Ku et al., 2016)
                  piwi
                  suppressible
                  Zip99C
                  (Smulders-Srinivasan and Lin, 2003)
                  Starting gene(s)
                  Interaction type
                  Interacting gene(s)
                  Reference
                  E2f1
                  suppressible
                  piwi
                  (Li and Lu, 2009)
                  Kr
                  enhanceable
                  piwi
                  (Gangaraju et al., 2011)
                  l(3)mbt
                  suppressible
                  piwi
                  (Janic et al., 2010)
                  RanGAP
                  enhanceable
                  piwi
                  (Gell and Reenan, 2013)
                  sd
                  suppressible
                  piwi
                  (Grimaud et al., 2006)
                  Su(var)3-3
                  suppressible
                  piwi
                  (Lepesant et al., 2020)
                  Tudor-SN
                  suppressible
                  piwi
                  (Ku et al., 2016)
                  yki
                  suppressible
                  piwi
                  (Zhang et al., 2015)
                  External Data
                  Subunit Structure (UniProtKB)
                  In the ovaries, part of a complex composed of at least Panx, nxf2, piwi and Nxt1 (PubMed:26472911, PubMed:26494711, PubMed:31368590, PubMed:31384064). The complex is knowns as Panx-induced co-transcriptional silencing (PICTS) complex, Panx-nxf2-dependent TAP/p15 silencing (Pandas complex), SFiNX (silencing factor interacting nuclear export variant) or piwi-Panx-nxf2-p15 (PPNP) complex (PubMed:26472911, PubMed:26494711, PubMed:31368590, PubMed:31384064). Interacts with vas; this interaction is RNA-independent (PubMed:16949822). Interacts with Dcr-1 and Fmr1; these interactions occur in polar granules (PubMed:16949822). Interacts (via N-terminal region) with CBX5 (via chromoshadow domain) (PubMed:17875665). Forms a complex with Hsp83 and Hop; probably Hop mediates the interaction between piwi and Hsp83 (PubMed:21186352). Forms a complex with Yb body components armi and fs(1)Yb; this interaction is required for proper piRNA loading and nuclear localization of piwi (PubMed:20966047). Interaction of Piwi and fs(1)Yb is likely to occur via armi (PubMed:20966047). Interacts (via the N-terminal region when unmethylated or symmetrically methylated at Arg-10) with papi (via Tudor domain) (PubMed:21447556, PubMed:29531043). Interacts with vret (PubMed:21831924). Interacts with Panx (PubMed:26472911, PubMed:26494711). Interacts with arx (PubMed:23913921, PubMed:23913922). Interacts with Tudor-SN (PubMed:26808625). Interacts with Nup358 (via N-terminus) (PubMed:29735528). Associates with the nuclear pore complex via interaction with Elys (PubMed:28472469). Interacts with thoc5; the interaction might be partly RNA-mediated (PubMed:28472469). Interacts with xmas-2 (PubMed:28472469).
                  (UniProt, Q9VKM1 )
                  Linkouts
                  BioGRID - A database of protein and genetic interactions.
                  DroID - A comprehensive database of gene and protein interactions.
                  MIST (protein-protein) - An integrated Molecular Interaction Database
                  Pathways
                  Signaling Pathways (FlyBase)
                  Metabolic Pathways
                  FlyBase
                  External Links
                  External Data
                  Linkouts
                  KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
                  Class of Gene
                  Genomic Location and Detailed Mapping Data
                  Chromosome (arm)
                  2L
                  Recombination map
                  2-44
                  Cytogenetic map
                  32C1-32C1
                  Sequence location
                  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

                  2-44

                  (FlyBase, 2016)

                  2-37.1

                  (Comeron et al., 2012)
                  Left of (cM)
                  Right of (cM)
                  Notes
                  Stocks and Reagents
                  Stocks (21)
                  Genomic Clones (21)
                  cDNA Clones (79)
                   

                  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 JBrowse for alignment of the cDNAs and ESTs to the gene model.

                  cDNA clones, fully sequenced
                  BDGP DGC clones
                  Other clones
                  Drosophila Genomics Resource Center cDNA clones

                  For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

                  cDNA Clones, End Sequenced (ESTs)
                  BDGP DGC clones
                  Other clones
                  RNAi and Array Information
                  Linkouts
                  DRSC - Results frm RNAi screens
                  Antibody Information
                  Laboratory Generated Antibodies
                   

                  polyclonal

                  (Tu et al., 2023, Zhang et al., 2018, Pandey et al., 2017, Vrettos et al., 2017, Wang et al., 2016, Sytnikova et al., 2014, Klattenhoff et al., 2009)

                  monoclonal

                  (Senti et al., 2015)
                  Commercially Available Antibodies
                   
                  Cell Line Information
                  Publicly Available Cell Lines
                   
                  Other Stable Cell Lines
                   
                  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)

                  The piwi product binds directly to repeat-associated small interfering RNAs (rasiRNAs).

                  (Vagin et al., 2006)

                  piwi is required for the accumulation of 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)
                  Relationship to Other Genes
                  Source for database merge of
                  Additional comments
                  Nomenclature History
                  Source for database identify of

                  Source for identity of: piwi CG6122

                  (Gong, 1999.11)
                  Nomenclature comments
                  Etymology
                  Synonyms and Secondary IDs (5)
                  Reported As
                  Symbol Synonym
                  CG6122
                  (Baumgartner et al., 2022, National Institute of Genetics Fly Stocks, 2022-, Na et al., 2016, Hosono et al., 2015, Ni et al., 2011.7.19, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Ni et al., 2008)
                  PIWI
                  (Glineburg and Nguyen, 2025, Krejčová and Bajgar, 2025, Pritam and Signor, 2025, Gamez et al., 2020, Meyer-Nava et al., 2020, Dou et al., 2019, Saha and Mishra, 2019, Azlan et al., 2016, Chen and Aravin, 2015, Wang et al., 2015, Eissenberg and Elgin, 2014, Ho et al., 2014, Sytnikova et al., 2014, Castel and Martienssen, 2013, Gomes et al., 2013, Dufourt et al., 2011, Kolaczkowski et al., 2011, Mendez et al., 2011, Lee and Langley, 2010, Klenov, 2008, Lin, 2008, Bai et al., 2007, Grewal and Elgin, 2007, Sofola et al., 2007, Zhou et al., 2007, Grimaud et al., 2006, Megosh et al., 2006, Rehwinkel et al., 2006, Morgan, 2002, Lowell, 2000)
                  Piwi
                  (Cacchione et al., 2025, Lee et al., 2025, Luan, 2025, Suyama and Kai, 2025, Zúniga-García and Riesgo-Escovar, 2025, Ho et al., 2024, Kotov et al., 2024, Rajeev et al., 2024, Yun et al., 2024, Lin et al., 2023, Parikh and Gangaraju, 2023, Santos et al., 2023, Tu et al., 2023, Arkov, 2022, Baumgartner et al., 2022, Benoit et al., 2022, He et al., 2022, Kotov et al., 2022, Rose et al., 2022, Eastwood et al., 2021, Fabry et al., 2021, Gong et al., 2021, Huang and Wong, 2021, Lauria Sneideman and Meller, 2021, Munafò et al., 2021, Onishi et al., 2021, Schneider and Imler, 2021, Vrettos et al., 2021, Wang and Lin, 2021, Wang et al., 2021, Adashev et al., 2020, Cacchione et al., 2020, Funk et al., 2020, Gonzalez et al., 2020, Kelleher et al., 2020, Lepesant et al., 2020, Mugat et al., 2020, Ninova et al., 2020, Ninova et al., 2020, Sankaranarayanan and Weil, 2020, Sato and Siomi, 2020, Shatskikh et al., 2020, Soleimani et al., 2020, Tindell et al., 2020, Tsuboyama et al., 2020, Yamaguchi et al., 2020, Yamashiro et al., 2020, Zhang and Cai, 2020, Batki et al., 2019, Fabry et al., 2019, Ge et al., 2019, Hirakata et al., 2019, Ishizu et al., 2019, Munafò et al., 2019, Murano et al., 2019, Russo et al., 2019, Schüpbach, 2019, Specchia et al., 2019, Zhao et al., 2019, Jankovics et al., 2018, Parikh et al., 2018, Rojas-Ríos and Simonelig, 2018, Sato and Siomi, 2018, Yamashiro and Siomi, 2018, Yashiro et al., 2018, Zhang et al., 2018, Dehghani and Lasko, 2017, Fefelova et al., 2017, Huang et al., 2017, Hyun, 2017, Ilyin et al., 2017, Karam et al., 2017, Luo and Lu, 2017, Pandey et al., 2017, Rogers et al., 2017, Ryazansky et al., 2017, Vrettos et al., 2017, Yang and Xi, 2017, Andress et al., 2016, Haase, 2016, Hayashi et al., 2016, Hirakata and Siomi, 2016, Jiang et al., 2016, Kelleher, 2016, Kotov et al., 2016, Ku et al., 2016, Lewis et al., 2016, Luo et al., 2016, Peng et al., 2016, Wang et al., 2016, Gonzalez et al., 2015, Han et al., 2015, Lin et al., 2015, Malone et al., 2015, Meller et al., 2015, Molla-Herman et al., 2015, Sato and Siomi, 2015, Sato et al., 2015, Hamada-Kawaguchi et al., 2014, Matzat and Lei, 2014, McElroy et al., 2014, Murota et al., 2014, Patil et al., 2014, Bausek, 2013, Czech et al., 2013, Darricarrère et al., 2013, Gomes et al., 2013, Guzzardo et al., 2013, Lasko, 2013, Le Thomas et al., 2013, Mani and Juliano, 2013, Ohtani et al., 2013, Olovnikov and Kalmykova, 2013, Peng and Lin, 2013, Rozhkov et al., 2013, Saito, 2013, Seervai and Wessel, 2013, Sentmanat et al., 2013, Vagin et al., 2013, Anand and Kai, 2012, Bozzetti et al., 2012, Kelleher et al., 2012, Kibanov et al., 2012, Kumar and Chen, 2012, McCue and Slotkin, 2012, Nishimasu et al., 2012, Olivieri et al., 2012, Preall et al., 2012, Stefanov et al., 2012, Vazquez-Pianzola and Suter, 2012, Eliazer and Buszczak, 2011, Ishizu et al., 2011, Khurana et al., 2011, Pane et al., 2011, Pek and Kai, 2011, Sato et al., 2011, Tiwari et al., 2011, Zhang et al., 2011, Kirino et al., 2010, Rigoutsos, 2010, Klattenhoff et al., 2009, Li et al., 2009, Miyoshi et al., 2009, Nishida et al., 2009, Czech et al., 2008, Kawamura et al., 2008, Lei et al., 2008, Shpiz et al., 2008, Antoniewski et al., 2007, Gunawardane et al., 2007, Gunawardane et al., 2007, Klenov et al., 2007, Lei, 2007, Lin, 2007, Nishida et al., 2007, Peng and Karpen, 2007, Prasanth and Spector, 2007, Saito et al., 2007, Yin and Lin, 2007, Zamore, 2007, Carthew, 2006, Dorner et al., 2006, Nystul and Spradling, 2006, Zamore, 2006, Gim et al., 2001)
                  piwi
                  (Alizada et al., 2025, Colmenares et al., 2025, Kina et al., 2025, Varoqui et al., 2025, Collins et al., 2024, Mancheno-Ferris et al., 2024, Patel et al., 2024, Sarkies, 2024, Yushkova, 2024, Asif-Laidin et al., 2023, Heigwer et al., 2023, Mohamed et al., 2023, Saha et al., 2023, Tang et al., 2023, Titlow et al., 2023, Voutyraki et al., 2023, Gorsi et al., 2022, Lim et al., 2022, National Institute of Genetics Fly Stocks, 2022-, Shapiro-Kulnane et al., 2022, Wang et al., 2022, Yang et al., 2022, Zinshteyn and Barbash, 2022, Casale et al., 2021, Gonzalez et al., 2021, Proshkina et al., 2021, Vidaurre and Chen, 2021, Yang, 2021, Zhang et al., 2021, Angelova et al., 2020, Duan et al., 2020, Ellison et al., 2020, Fort-Aznar et al., 2020, Hinnant et al., 2020, Kordyukova et al., 2020, Mérel et al., 2020, Onishi et al., 2020, Parker et al., 2020, Shi et al., 2020, Chang et al., 2019, Kina et al., 2019, Kneuss et al., 2019, Kolliopoulou et al., 2019, Kotov et al., 2019, Molnar et al., 2019, Park et al., 2019, Schaub et al., 2019, Stein et al., 2019, Trcek and Lehmann, 2019, Wakisaka et al., 2019, Whittle and Extavour, 2019, Yang et al., 2019, Gene Disruption Project members, 2018-, Hao et al., 2018, Huang et al., 2018, Hughes et al., 2018, Kang et al., 2018, Kordyukova et al., 2018, Levis, 2018.8.30, Ma et al., 2018, Mehrotra and Deshpande, 2018, Sun et al., 2018, Tasnim and Kelleher, 2018, Théron et al., 2018, Tian and Wang, 2018, van den Beek et al., 2018, Akkouche et al., 2017, Marie et al., 2017, Rashpa et al., 2017, Sousa-Victor et al., 2017, Specchia et al., 2017, Teixeira et al., 2017, Abramov et al., 2016, Basu et al., 2016, Blumenstiel et al., 2016, Fagegaltier et al., 2016, Klein et al., 2016, Na et al., 2016, Reid and O'Brochta, 2016, Wang et al., 2016, Flores et al., 2015, Ishizu et al., 2015, Mikhaleva et al., 2015, Mugat et al., 2015, Onorati et al., 2015, Sato et al., 2015, Senti et al., 2015, Sienski et al., 2015, Yu et al., 2015, Zhang et al., 2015, Atikukke et al., 2014, Basquin et al., 2014, Blanchard et al., 2014, Dufourt et al., 2014, Hamada-Kawaguchi et al., 2014, Hayashi et al., 2014, Le Thomas et al., 2014, Lim et al., 2014, Ma et al., 2014, Malone et al., 2014, Mani et al., 2014, Minakhina et al., 2014, Pargett et al., 2014, Ren et al., 2014, Ross et al., 2014, Wang et al., 2014, Bausek, 2013, Czech et al., 2013, Darricarrère et al., 2013, Dennis et al., 2013, Dönertas et al., 2013, Gell and Reenan, 2013, Handler et al., 2013, Huang et al., 2013, Jin et al., 2013, Li et al., 2013, Muerdter et al., 2013, Ohtani et al., 2013, Bozzetti et al., 2012, Japanese National Institute of Genetics, 2012.5.21, McEachern, 2012, Meier et al., 2012, Menon and Meller, 2012, Siddiqui et al., 2012, Sienski et al., 2012, Simmons et al., 2012, Zhang et al., 2012, Castillo et al., 2011, Ding and Lu, 2011, Eliazer and Buszczak, 2011, Gangaraju et al., 2011, Handler et al., 2011, Klenov et al., 2011, Klenov et al., 2011, Liu et al., 2011, Miles et al., 2011, Ni et al., 2011, Parrott et al., 2011, Pek and Kai, 2011, Pek and Kai, 2011, Poulton et al., 2011, Qi et al., 2011, Shpiz et al., 2011, Tchurikov and Kretova, 2011, Wang and Elgin, 2011, Zamparini et al., 2011, Fernández-Ayala et al., 2010, Haase et al., 2010, Janic et al., 2010, Lu and Clark, 2010, Moshkovich and Lei, 2010, Nagao et al., 2010, Olivieri et al., 2010, Rouget et al., 2010, Saito et al., 2010, Simmons et al., 2010, Smulders-Srinivasan et al., 2010, Stefanov et al., 2010, Thomsen et al., 2010, Belinco et al., 2009, Chau et al., 2009, Habayeb et al., 2009, Jepson and Reenan, 2009, Kavi and Birchler, 2009, Kirino et al., 2009, Lau et al., 2009, Li and Lu, 2009, Malone et al., 2009, Robine et al., 2009, Saito et al., 2009, Shpiz et al., 2009, Szakmary et al., 2009, Venken et al., 2009, Berdnik et al., 2008, Blumenstiel et al., 2008, Chambeyron et al., 2008, Chau et al., 2008, Chotkowski et al., 2008, Cross and Simmons, 2008, Desset et al., 2008, Kalmykova et al., 2008, Kavi and Birchler, 2008, McGraw et al., 2008, Ronsseray et al., 2008, Bhadra et al., 2007, Brennecke et al., 2007, Brower-Toland et al., 2007, Georlette et al., 2007, Glaser et al., 2007, Josse et al., 2007, Kavi and Birchler, 2007, Kruger, 2007, Pelisson et al., 2007, Ronsseray et al., 2007, Shpiz et al., 2007, Simmons et al., 2007, Sofola et al., 2007, Wu and Silverman, 2007, Xing et al., 2007, Yang et al., 2007, Zaratiegui, 2007, Christensen and Cook, 2006.8.30, Haynes et al., 2006, Kavi et al., 2006, Lei and Corces, 2006, Lei and Corces, 2006, Megosh et al., 2006, Musters et al., 2006, Payne and Braun, 2006, Saito et al., 2006, Shigenobu et al., 2006, Vagin et al., 2006, Vagin et al., 2006, Kalmykova et al., 2005, Xie et al., 2005, Gonzalez-Reyes, 2003)
                  Name Synonyms
                  P-element induced wimpy testis
                  (Pek and Kai, 2011, Lin and Spradling, 1997)
                  PIWI
                  (Ronsseray et al., 2007, Seydoux and Braun, 2006)
                  Piwi
                  (Hirakata and Siomi, 2016, Rozhkov et al., 2013, Simkin et al., 2013, Carmi, 2006)
                  piwi
                  (Gell and Reenan, 2013, Kaczmarczyk and Kopp, 2011, Saito et al., 2009, Cross and Simmons, 2008, Josse et al., 2007, Xing et al., 2007, Kavi et al., 2006)
                  Secondary FlyBase IDs
                    Datasets (3)
                    Study focus (3)
                    Experimental Role
                    Project
                    Project Type
                    Title
                    • bait_protein
                    modENCODE_chromatin_proteins_cells
                    Genome-wide localization of chromosomal proteins in cell lines by ChIP-chip and ChIP-Seq.
                    • allele_used
                    modENCODE_chromatin_proteins_flies
                    Genome-wide localization of chromosomal proteins in fly tissues by ChIP-chip and ChIP-Seq.
                    • allele_used
                    modENCODE_chromatin_histones_flies
                    Genome-wide localization of histones and their modifications in fly tissues by ChIP-chip and ChIP-Seq.
                    Study result (0)
                    Result
                    Result Type
                    Title
                    External Crossreferences and Linkouts ( 48 )
                    Sequence Crossreferences
                    NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
                    GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
                    GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
                    RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
                    UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
                    UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
                    UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
                    Other crossreferences
                    AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
                    BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
                    DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
                    EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
                    FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
                    FlyMine - An integrated database for Drosophila genomics
                    InterPro - A database of protein families, domains and functional sites
                    KEGG Genes - Molecular building blocks of life in the genomic space.
                    MARRVEL_MODEL - MARRVEL (model organism gene)
                    PDB - An information portal to biological macromolecular structures
                    Linkouts
                    BioGRID - A database of protein and genetic interactions.
                    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
                    DroID - A comprehensive database of gene and protein interactions.
                    DRSC - Results frm RNAi screens
                    Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
                    FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
                    FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                    Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
                    Flygut - An atlas of the Drosophila adult midgut
                    Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                    KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
                    MIST (protein-protein) - An integrated Molecular Interaction Database
                    References (605)