FlyBase Gene Report: Dmel\tws

General Information

Symbol

Dmel\tws

Species

D. melanogaster

Name

twins

Annotation Symbol

CG6235

Feature Type

protein_coding_gene

FlyBase ID

FBgn0004889

Gene Model Status

Current

Stock Availability

30 publicly available

Gene Summary

twins (tws) encodes a regulatory subunit of protein phosphatase 2A (PP2A) and is involved in many developmental processes and signaling pathways. [Date last reviewed: 2018-09-13] (FlyBase Gene Snapshot)

All Summaries

Gene Snapshot
Alliance
Auto summary
Pathway
Gene Group
UniProtKB
Interactive Fly

Also Known As

aar, PP2A, PR55, phosphoprotein phosphatase 2A 55 kDa, l(3)02414

Key Links

Genomic Location

Cytogenetic map

85F13-85F14

Sequence location

3R:10,126,129..10,139,772 [-]

Recombination map

3-49

RefSeq locus

NT_033777 REGION:10126129..10139772

Sequence

Get Decorated FASTA

Genomic Maps

JBrowse

Other Genome Views

The following external sites may use different assemblies or annotations than FlyBase.

Function

Gene Ontology (GO) Annotations (17 terms)

Molecular Function (1 term)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

enables protein phosphatase regulator activity

inferred from mutant phenotype

(Mayer-Jaekel et al., 1994)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

enables protein phosphatase regulator activity

inferred from electronic annotation with InterPro:IPR000009

(InterPro Project Members, 2004-)

inferred from biological aspect of ancestor with PANTHER:PTN000221745

(GO Reference Genome Project, 2011-)

inferred from sequence or structural similarity with UniProtKB:P63151

(Gene Ontology Curators, 2002-)

Biological Process (13 terms)

Terms Based on Experimental Evidence (13 terms)

CV Term

Evidence

References

involved_in centrosome cycle

inferred from mutant phenotype

(Dobbelaere et al., 2008)

involved_in imaginal disc pattern formation

inferred from mutant phenotype

(Uemura et al., 1993)

involved_in maintenance of neuroblast polarity

inferred from mutant phenotype

(Chabu and Doe, 2009)

involved_in metaphase/anaphase transition of mitotic cell cycle

inferred from mutant phenotype

(Mayer-Jaekel et al., 1993)

involved_in mitotic cell cycle

inferred from mutant phenotype

(Chen et al., 2007)

involved_in negative regulation of neuroblast proliferation

inferred from mutant phenotype

(Chabu and Doe, 2009)

involved_in positive regulation of canonical Wnt signaling pathway

inferred from mutant phenotype

(Bajpai et al., 2004)

inferred from genetic interaction with FLYBASE:dsh; FB:FBgn0000499

(Bajpai et al., 2004)

inferred from genetic interaction with FLYBASE:sgg; FB:FBgn0003371

(Bajpai et al., 2004)

inferred from genetic interaction with FLYBASE:pan; FB:FBgn0085432

(Bajpai et al., 2004)

involved_in positive regulation of smoothened signaling pathway

inferred from mutant phenotype

(Su et al., 2011)

involved_in positive regulation of Toll signaling pathway

inferred from genetic interaction with FLYBASE:Gyc76C; FB:FBgn0266136

(Kanoh et al., 2021)

involved_in protein stabilization

inferred from direct assay

(Brownlee et al., 2011)

inferred from mutant phenotype

(Brownlee et al., 2011)

involved_in regulation of centriole replication

inferred from mutant phenotype

(Brownlee et al., 2011)

involved_in regulation of mitotic metaphase/anaphase transition

inferred from mutant phenotype

(Mayer-Jaekel et al., 1993)

involved_in sensory organ development

inferred from mutant phenotype

(Shiomi et al., 1994)

Terms Based on Predictions or Assertions (0 terms)

Cellular Component (3 terms)

Terms Based on Experimental Evidence (2 terms)

CV Term

Evidence

References

located_in cytosol

inferred from direct assay

(Wang et al., 2011, Chabu and Doe, 2009, Dobbelaere et al., 2008, Mayer-Jaekel et al., 1994)

inferred from high throughput direct assay

(Lye et al., 2014)

located_in nucleus

inferred from direct assay

(Chabu and Doe, 2009)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

is_active_in cytosol

inferred from biological aspect of ancestor with PANTHER:PTN000221745

(GO Reference Genome Project, 2011-)

part_of protein phosphatase type 2A complex

inferred from biological aspect of ancestor with PANTHER:PTN000221745

(GO Reference Genome Project, 2011-)

inferred from electronic annotation with InterPro:IPR000009

(InterPro Project Members, 2004-)

inferred from sequence or structural similarity with UniProtKB:P63151

(Gene Ontology Curators, 2002-)

Gene Group (FlyBase)

TWS-PROTEIN PHOSOPHATASE 2A COMPLEX

Pathway (FlyBase)

POSITIVE REGULATORS OF WNT-TCF SIGNALING PATHWAY

POSITIVE REGULATORS OF HEDGEHOG SIGNALING PATHWAY

POSITIVE REGULATORS OF TOLL-NF-kappaB SIGNALING PATHWAY

Protein Family (UniProt)

Belongs to the phosphatase 2A regulatory subunit B family. (P36872)

Protein Signatures (InterPro)

Summaries

Gene Snapshot

twins (tws) encodes a regulatory subunit of protein phosphatase 2A (PP2A) and is involved in many developmental processes and signaling pathways. [Date last reviewed: 2018-09-13]

Pathway (FlyBase)

POSITIVE REGULATORS OF WNT-TCF SIGNALING PATHWAY -: Positive regulators of Wnt-TCF (canonical Wnt) signaling up-regulate the pathway, resulting in the enhancement of transcriptional regulation mediated by β-catenin (arm).
POSITIVE REGULATORS OF HEDGEHOG SIGNALING PATHWAY -: Positive regulators of hedgehog signaling up-regulate the pathway, resulting in the activation of transcription of hh-responsive genes.
POSITIVE REGULATORS OF TOLL-NF-kappaB SIGNALING PATHWAY -: In Drosophila, the canonical Toll signaling pathway is initiated by the binding of a spatzle ligand to Toll (Tl) or a Toll-like receptor leading to the nuclear localization of the NF-κB (dl or Dif) transcription factor. Positive regulators of Toll-NF-κB Signaling lead to the increased nuclear accumulation of the NF-κB proteins and regulated expression of target genes. (Adapted from FBrf0091014 and FBrf0223077).

Gene Group (FlyBase)

TWS-PROTEIN PHOSOPHATASE 2A COMPLEX -: tws-PP2A is a serine/threonine phosphatase complex characterized by the presence of a tws regulatory B subunit. (Adapted from FBrf0232706.)

Protein Function (UniProtKB)

Could perform a substrate recognition function or could be responsible for targeting the enzyme complex to the appropriate subcellular compartment.

(UniProt, P36872)

Summary (Interactive Fly)

Protein phosphatase 2A - B subunit - mutation causes a peripheral nervous system defect similar to those of numb and musashi - mutation also causes a pattern duplication in imaginal discs

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\tws 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 P36872)

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

Comments on Gene Model

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

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

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

Gene model reviewed during 5.46

Gene model reviewed during 5.55

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

tws-RA

FBtr0082186

NM_057532

2520

499

tws-RB

FBtr0082188

NM_134286

2896

443

tws-RC

FBtr0082191

NM_134285

2242

443

tws-RD

FBtr0082189

NM_134284

3086

443

tws-RE

FBtr0082190

NM_057533

3819

443

tws-RF

FBtr0082187

NM_169329

3664

499

tws-RG

FBtr0082192

NM_169330

2400

443

tws-RH

FBtr0082193

NM_169331

2300

443

tws-RI

FBtr0345177

NM_001300340

3856

499

Additional Transcript Data and Comments

Reported size (kB)

5.0, 3.8-4.0, 3.7, 2.75, 2.5 (northern blot)

(Mayer-Jaekel et al., 1993)

Comments

External Data

Crossreferences

Polypeptide Data

Annotated Polypeptides

Name

FlyBase ID

Predicted MW (kDa)

Length (aa)

Theoretical pI

UniProt

RefSeq ID

GenBank

tws-PA

FBpp0081664

57.0

499

7.10

FBpp0081666

50.8

443

6.38

FBpp0081669

50.8

443

6.38

FBpp0081667

50.8

443

6.38

FBpp0081668

50.8

443

6.38

FBpp0081665

57.0

499

7.10

FBpp0081670

50.8

443

6.38

FBpp0081671

50.8

443

6.38

FBpp0311386

57.0

499

7.10

P36872

NP_001287269

AHN57268

Polypeptides with Identical Sequences

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

499 aa isoforms: tws-PA, tws-PF, tws-PI

443 aa isoforms: tws-PB, tws-PC, tws-PD, tws-PE, tws-PG, tws-PH

Additional Polypeptide Data and Comments

Reported size (kDa)

456, 443 (aa); 52, 51 (kD)

(Mayer-Jaekel et al., 1993)

443 (aa); 51 (kD predicted)

(Uemura et al., 1993)

Comments

External Data

Subunit Structure (UniProtKB)

PP2A exists in several trimeric forms, all of which consist of a core composed of a catalytic subunit associated with a 65 kDa regulatory subunit (PR65) (subunit A). The core complex associates with a third, variable subunit (subunit B), which confers distinct properties to the holoenzyme.

(UniProt, P36872)

Crossreferences

InterPro - A database of protein families, domains and functional sites

Linkouts

Sequences Consistent with the Gene Model

Nucleotide / Polypeptide Records

UniProt

Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\tws 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).

-0.39

Transcript Expression

No Assay Recorded

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 4

(Mayer-Jaekel et al., 1993)

larval stage

adult optic lobe

(Mayer-Jaekel et al., 1993)

pupal stage & spermatogenesis

testis

(Mayer-Jaekel et al., 1993)

adult stage & oogenesis

oocyte

(Mayer-Jaekel et al., 1993)

adult stage oogenesis stage S6 -- S14

nurse cell

(Mayer-Jaekel et al., 1993)

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 3 -- 4

organism | ubiquitous

(Mayer-Jaekel et al., 1993)

embryonic stage 10 -- 17

gonad

(Mayer-Jaekel et al., 1993)

neuroblast

(Mayer-Jaekel et al., 1993)

embryonic stage 17

Malpighian tubule

(Mayer-Jaekel et al., 1993)

anal pad

(Mayer-Jaekel et al., 1993)

presumptive embryonic/larval central nervous system

(Mayer-Jaekel et al., 1993)

embryonic/larval hindgut

(Mayer-Jaekel et al., 1993)

larval stage

adult optic lobe

(Mayer-Jaekel et al., 1993)

pharate adult stage

testis

(Mayer-Jaekel et al., 1993)

adult stage & oogenesis

oocyte

(Mayer-Jaekel et al., 1993)

adult stage oogenesis stage S6 -- S14

nurse cell

(Mayer-Jaekel et al., 1993)

northern blot

Stage

Tissue/Position (including subcellular localization)

Reference

larval stage

eye-antennal disc

(Uemura et al., 1993)

leg disc

(Uemura et al., 1993)

larval central nervous system

(Uemura et al., 1993)

wing disc

(Uemura et al., 1993)

third instar larval stage

(Uemura et al., 1993)

imaginal disc

(Mayer-Jaekel et al., 1993)

Additional Descriptive Data

tws transcripts are found at high levels in ovarioles. They are first detected at stage S6 in nurse cells and are later transported to the egg cell. In embryos, they are present at high levels at the syncytial blastoderm stage and then decrease. They are later detected in the migrating gonads, CNS, anal pads, Malpighian tubules, and hindgut. In larvae, transcripts are detected in regions of cell proliferation in the optic lobes. In pupae, they are found in a uniform distribution in imaginal discs and testes.

(Mayer-Jaekel et al., 1993)

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

immunolocalization

Stage

Tissue/Position (including subcellular localization)

Reference

larval stage

wing disc

(Bajpai et al., 2004)

western blot

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage & larval stage & pupal stage

(Uemura et al., 1993)

larval stage

larval central nervous system

(Uemura et al., 1993)

imaginal disc

(Uemura et al., 1993)

embryonic/larval salivary gland

(Uemura et al., 1993)

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in cytosol

inferred from direct assay

(Wang et al., 2011, Chabu and Doe, 2009, Dobbelaere et al., 2008, Mayer-Jaekel et al., 1994)

inferred from high throughput direct assay

(Lye et al., 2014)

located_in nucleus

inferred from direct assay

(Chabu and Doe, 2009)

Expression Deduced from Reporters

Reporter: Mi{PT-GFSTF.1}tws^{MI09216-GFSTF.1}

Stage

Tissue/Position (including subcellular localization)

Reference

third instar larval stage

larval brain

(Nagarkar-Jaiswal et al., 2015)

Reporter: PBac{681.P.FSVS-1}tws^CPTI001600

Stage

Tissue/Position (including subcellular localization)

Reference

adult stage

medulla

Comment: medulla - photoreceptors?

(Knowles-Barley, 2011.8.24)

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\tws in JBrowse

RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region

View exonic expression by developmental stage for Dmel\tws

View exonic expression by tissue for Dmel\tws

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

FBgn0004889

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

47877/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0004889

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0004889

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0004889

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG6235

Flygut - An atlas of the Drosophila adult midgut

FBgn0004889

Images

FlyBase Wiki

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 103 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 23 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of tws

Transgenic constructs containing regulatory region of tws

Aberrations (Deficiencies and Duplications) ( 10 )

Inferred from experimentation ( 10 )

Gene disrupted in

Df(3R)BSC38

(Andrade et al., 2003.7.4)

Df(3R)BSC526

(Christensen et al., 2008.6.11)

Df(3R)BSC621

(Cook et al., 2008.9.29)

Df(3R)by10

(BDGP Project Members, 1994-1999)

Df(3R)by62

(Merigliano et al., 2017, Spradling et al., 1999, Szeged Stock Center, 1998-, Deak et al., 1997, BDGP Project Members, 1994-1999, Uemura et al., 1993)

Df(3R)Exel6265

(Ryder, 2004.4.26)

Tp(3;2)by62

(BDGP Project Members, 1994-1999)

Gene not disrupted in

Df(3L)Cat

(Szeged Stock Center, 1998-)

Df(3L)W10

(Szeged Stock Center, 1998-)

Df(3R)23D1

(Szeged Stock Center, 1998-)

Df(3R)by10

(Spradling et al., 1999)

Inferred from location ( 5 )

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

lethal, with Scer\GAL4^tim.PE

tws^UAS.cGa

lethal, with Scer\GAL4^αTub84B.PL

tws^GL00670

lethal (with Df(3R)Exel6265)

tws^{MI09216-TG4.1}

lethal | adult stage | recessive

tws^aar-1

lethal | embryonic stage | maternal effect

tws⁰²⁴¹⁴

lethal | larval stage | recessive

lethal | pharate adult stage | recessive

tws^aar-1

lethal | pupal stage | recessive

lethal - all die before end of larval stage

tws⁴³⁰

lethal - all die before end of prepupal stage | recessive

lethal - all die before end of P-stage | recessive

tws^S061805

lethal - all die before end of pupal stage

lethal - all die before end of pupal stage | recessive

lethal - all die during embryonic stage

tws⁰²⁴¹⁴

partially lethal, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

tws^GD10742

partially lethal - majority die

tws^v158

partially lethal - majority die (with Df(3R)by62)

tws^S134601

partially lethal - majority die | recessive

viable

viable, with Scer\GAL4^Mef2.PR

tws^GD10742

viable, with Scer\GAL4^pnr-MD237

tws^GD10742

viable, with Scer\GAL4^tin.CΔ4

tws^GD10742

viable, with Scer\GAL4^{VP16.nanos.UTR}

tws^UAS.GFP

Sterility

Allele

female semi-sterile, with Scer\GAL4^{VP16.nanos.UTR}, Scer\GAL4^VP16.otu, Scer\GAL4^nanos.PG

tws^HMS01578

female sterile, with Scer\GAL4^{VP16.mat.αTub67C}

tws^GL00670

female sterile | recessive

tws^aar-2

sterile, with Scer\GAL4^{VP16.mat.αTub67C}

tws^GL00670

Other Phenotypes

Allele

abnormal behavior | adult stage

tws^CB-5218-3

abnormal behavior | recessive

abnormal body color, with Scer\GAL4^pnr-MD237

tws^GD10742

abnormal cell shape | somatic clone

tws^j11C8

abnormal locomotor rhythm, with Scer\GAL4^P2.4.Pdf

tws^UAS.cGa

abnormal meiotic cell cycle

tws^v158

abnormal mitotic cell cycle

tws^v158

abnormal mitotic cell cycle | larval stage

tws^aar-1

abnormal mitotic cell cycle | third instar larval stage

abnormal neuroanatomy, with Scer\GAL4⁴⁷⁷, Scer\GAL4^ppk.PU

tws^HMS04498

abnormal neuroanatomy | adult stage | progressive, with Scer\GAL4^elav.PU

tws^UAS.cBa

decreased cell size (with Df(3R)Exel6265)

tws^j11C8

increased cell death | embryonic stage, with Scer\GAL4^Act5C.PU

tws^UAS.cBa

increased cell death | embryonic stage, with Scer\GAL4^elav.PU

tws^UAS.cBa

increased cell number (with Df(3R)Exel6265)

tws^j11C8

increased cell number | somatic clone

tws^j11C8

increased occurrence of cell division | larval stage

tws^aar-1

radiation sensitive | third instar larval stage

tws⁴³⁰

radiation sensitive | third instar larval stage (with Df(3R)by62)

radiation sensitive | third instar larval stage (with tws¹⁹⁶)

tws^j11C8

radiation sensitive | third instar larval stage (with tws^j11C8)

tws¹⁹⁶

short lived | heat sensitive, with Scer\GAL4^elav.PU

tws^UAS.cBa

some die during embryonic stage

tws⁰²⁴¹⁴

some die during immature adult stage | recessive

some die during pharate adult stage | recessive

tws^S061805

some die during prepupal stage | recessive

some die during pupal stage

some die during pupal stage | recessive

uncoordinated

visible, with Scer\GAL4^pnr-MD237

tws^GD10742

wild-type

tws^+tMa

Phenotype manifest in

Allele

abdomen | maternal effect

tws⁰²⁴¹⁴

adult brain | progressive, with Scer\GAL4^elav.PU

tws^UAS.cBa

chromatin & mitotic anaphase

tws^aar-1

chromosome, centromeric region

tws^v158

dendrite, with Scer\GAL4⁴⁷⁷, Scer\GAL4^ppk.PU

tws^HMS04498

embryo | maternal effect, with Scer\GAL4^{VP16.mat.αTub67C}

tws^GL00670

embryonic/larval optic lobe | somatic clone

tws^j11C8

eo neuron

eye

ganglion | larval stage

tws^v158

germarium

tws^v158

haltere disc

larval brain & nuclear chromosome

tws^v158

larval brain | third instar larval stage

tws⁴³⁰

larval brain | third instar larval stage (with Df(3R)by62)

larval brain | third instar larval stage (with tws¹⁹⁶)

larval brain | third instar larval stage (with tws⁴³⁰)

larval brain | third instar larval stage (with tws^j11C8)

larval neuroblast | somatic clone | increased number

tws^j11C8

larval neuroblast | somatic clone | increased number (with Df(3R)Exel6265)

tws^j11C8

macrochaeta

mesonotum, with Scer\GAL4^pnr-MD237

tws^GD10742

microchaeta

mitochondrion

tws^j11C8

mitochondrion, with Scer\GAL4^elav.PU

mitochondrion, with Scer\GAL4^en.PU

mitotic anaphase

tws^v158

mitotic anaphase & neuroblast

tws^S061805

mitotic cell cycle

tws^v158

mitotic metaphase & neuroblast

tws^S061805

mitotic spindle | third instar larval stage

tws^aar-1

multidendritic neuron, with Scer\GAL4⁴⁷⁷, Scer\GAL4^ppk.PU

tws^HMS04498

neuroblast

tws⁶⁰

neuroblast (with tws⁶⁰)

tws⁰²⁴¹⁴

neuroblast (with tws⁰²⁴¹⁴)

tws⁶⁰

neuroblast | larval stage

tws^aar-1

nuclear chromosome & mitotic anaphase

tws^aar-1

nuclear chromosome | third instar larval stage

tws⁴³⁰

nuclear chromosome | third instar larval stage (with Df(3R)by62)

nuclear chromosome | third instar larval stage (with tws¹⁹⁶)

nuclear chromosome | third instar larval stage (with tws⁴³⁰)

nuclear chromosome | third instar larval stage (with tws^j11C8)

nucleus | third instar larval stage

tws⁴³⁰

nucleus | third instar larval stage (with Df(3R)by62)

nucleus | third instar larval stage (with tws¹⁹⁶)

tws^j11C8

nucleus | third instar larval stage (with tws^j11C8)

tws¹⁹⁶

ovary

tws^v158

spindle & neuroblast

tws^S061805

testis

tws^v158

thecogen cell

tormogen cell

trichogen cell

trichogen cell | increased number, with Scer\GAL4^pnr-MD237

tws^GD10742

trichome of the posterior wing margin

wing disc | larval stage

wing disc posterior compartment

wing margin | somatic clone

tws^j11C8

wing margin bristle | ectopic, with Scer\GAL4^hs.PB, tws⁶⁰

tws^UAS.cBa

wing margin bristle | ectopic, with Scer\GAL4^hs.PB, tws^UAS.cBa

tws⁶⁰

Orthologs

Downloads

Download All DIOPT Orthologs

Human Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Homo sapiens (Human) (10)

Hsap\PPP2R2D

14 of 14

Yes

Hsap\PPP2R2A

13 of 14

Yes

Hsap\PPP2R2B

13 of 14

Yes

Hsap\PPP2R2C

13 of 14

Yes

Hsap\DDB2

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Rattus norvegicus (Norway rat) (10)

Rnor\Ppp2r2d

14 of 14

Yes

Rnor\Ppp2r2a

13 of 14

Yes

Rnor\Ppp2r2b

13 of 14

Yes

Rnor\Ppp2r2c

11 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Mus musculus (laboratory mouse) (10)

Mmus\Ppp2r2d

14 of 14

Yes

Mmus\Ppp2r2a

13 of 14

Yes

Mmus\Ppp2r2b

13 of 14

Yes

Mmus\Ppp2r2c

12 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Xenopus tropicalis (Western clawed frog) (11)

Xtro\ppp2r2d

11 of 13

Yes

Xtro\ppp2r2b

10 of 13

Yes

Xtro\ppp2r2a

7 of 13

Yes

Xtro\ppp2r2c

7 of 13

Yes

Xtro\adss1

1 of 13

Xtro\gnb5

1 of 13

Xtro\lrp1b

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

Danio rerio (Zebrafish) (12)

Drer\ppp2r2ab

13 of 14

Yes

Drer\ppp2r2bb

13 of 14

Yes

Drer\ppp2r2d

12 of 14

Yes

Drer\ppp2r2aa

10 of 14

Yes

Drer\ppp2r2ca

8 of 14

Yes

Drer\ppp2r2cb

8 of 14

Yes

Drer\ppp2r2ba

3 of 14

Yes

Drer\gnb4b

1 of 14

Drer\poc1a

1 of 14

Drer\poc1b

1 of 14

Drer\si:cabz01066312.1

1 of 14

Drer\wdr5

1 of 14

Caenorhabditis elegans (Nematode, roundworm) (4)

Cele\sur-6

14 of 14

Yes

Cele\gpb-1

1 of 14

Cele\prp-19

1 of 14

Cele\wdr-37

1 of 14

Anopheles gambiae (African malaria mosquito) (6)

Agam\AgaP_AGAP004502

12 of 12

Yes

1 of 12

1 of 12

1 of 12

1 of 12

Yes

Agam\AgaP_AGAP012731

1 of 12

Arabidopsis thaliana (thale-cress) (6)

Atha\ATB BETA

12 of 13

Yes

Atha\B ALPHA

12 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

Yes

Saccharomyces cerevisiae (Brewer's yeast) (1)

Scer\CDC55

13 of 13

Yes

Schizosaccharomyces pombe (Fission yeast) (3)

Spom\pab1

11 of 12

Yes

Spom\SPAC227.12

1 of 12

Spom\SPBC713.05

1 of 12

Escherichia coli (enterobacterium) (0)

Other Organism Orthologs (via OrthoDB)

Data provided directly from OrthoDB:tws. Refer to their site for version information.

Paralogs

Downloads

Download All DIOPT Paralogs

Paralogs (via DIOPT v9.1)

Gene Symbol

Score

Source

Alignment

Drosophila melanogaster (Fruit fly) (3)

CG7568

1 of 13

Gβ13F

1 of 13

wds

1 of 13

Human Disease Associations

FlyBase Human Disease Model Reports

spinocerebellar ataxia 12

Disease Ontology (DO) Annotations

Models Based on Experimental Evidence ( 1 )

Allele

Disease

Evidence

References

tws^UAS.cBa

model of autosomal dominant cerebellar ataxia

CEA

(Wang et al., 2011)

Potential Models Based on Orthology ( 1 )

Human Ortholog

Disease

Evidence

References

PPP2R2B; protein phosphatase 2 regulatory subunit Bbeta

model of spinocerebellar ataxia 12

IEA

(FlyBase, 2019-)

Modifiers Based on Experimental Evidence ( 1 )

Allele

Disease

Interaction

References

tws^UAS.cBa

model of autosomal dominant cerebellar ataxia

is ameliorated by Sod2^UAS.cMa

(Wang et al., 2011)

ameliorates tauopathy

modeled by Hsap\MAPT^UAS.cWa

(Yeh et al., 2010)

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?

PPP2R2D; protein phosphatase 2 regulatory subunit Bdelta

14 of 14

613992

PPP2R2A; protein phosphatase 2 regulatory subunit Balpha

13 of 14

604941

PPP2R2B; protein phosphatase 2 regulatory subunit Bbeta

13 of 14

PROTEIN PHOSPHATASE 2, REGULATORY SUBUNIT B, BETA; PPP2R2B

SPINOCEREBELLAR ATAXIA 12; SCA12

spinocerebellar ataxia 12

PPP2R2C; protein phosphatase 2 regulatory subunit Bgamma

13 of 14

605997

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

tws - CG33229

experimental knowledge based

(Guruharsha et al., 2011)

tws - HeT-A\gag

pull down, Identification by mass spectrometry

(Morgunova et al., 2021)

tws - Lam

anti tag coimmunoprecipitation, anti tag western blot

(Mehsen et al., 2018)

tws - Nup107

anti tag coimmunoprecipitation, anti tag western blot

(Mehsen et al., 2018)

tws - Pp2A-29B

pull down, western blot

(Larouche et al., 2021)

tws - S6k

anti tag coimmunoprecipitation, Identification by mass spectrometry

(Vinayagam et al., 2016)

tws - aPKC

anti bait coimmunoprecipitation, western blot

(Chabu and Doe, 2009)

tws - baf

proximity ligation assay, fluorescence microscopy, anti tag coimmunoprecipitation, anti tag western blot

(Mehsen et al., 2018)

tws - baz

anti tag coimmunoprecipitation, anti tag western blot

(Krahn et al., 2009)

tws - chico

anti tag coimmunoprecipitation, Identification by mass spectrometry

(Vinayagam et al., 2016)

tws - ci

anti tag coimmunoprecipitation, western blot

(Su et al., 2011)

tws - endos

pull down, western blot, enzymatic study, ultraviolet-visible spectroscopy, anti tag coimmunoprecipitation, anti tag western blot, coimmunoprecipitation, autoradiography

(Larouche et al., 2021, Kim et al., 2012)

tws - fd59A

anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based

(Rhee et al., 2014)

tws - gwl

proximity ligation assay, fluorescence microscopy, anti tag coimmunoprecipitation, anti tag western blot

(Wang et al., 2016)

tws - mei-S332

anti tag coimmunoprecipitation, peptide massfingerprinting

(Chen et al., 2007)

tws - mts

coimmunoprecipitation, western blot, pull down, anti tag coimmunoprecipitation, anti tag western blot

(Larouche et al., 2021, Kim et al., 2019, Mehsen et al., 2018)

tws - vtd

anti tag coimmunoprecipitation, anti tag western blot

(Kim et al., 2019)

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

tws

suppressible

aPKC

(Chabu and Doe, 2009)

tws

suppressible

arm

(Bajpai et al., 2004)

tws

suppressible

Cdc27

(Deak et al., 2003)

tws|polo

suppressible

endos

(Rangone et al., 2011)

tws

suppressible

grp

(Merigliano et al., 2017)

tws

suppressible

Irbp

(Merigliano et al., 2017)

tws

suppressible

mei-9

(Merigliano et al., 2017)

tws

suppressible

nbs

(Merigliano et al., 2017)

tws

suppressible

Sod2

(Wang et al., 2011)

tws

suppressible

spn-A

(Merigliano et al., 2017)

tws

suppressible

tefu

(Merigliano et al., 2017)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

arm

enhanceable

tws

(Greaves et al., 1999)

Cdc27

enhanceable

tws

(Deak et al., 2003)

dom

enhanceable

tws

(Kwon et al., 2013)

dsh

suppressible

tws

(Bajpai et al., 2004)

Egfr

enhanceable

tws

(Read et al., 2005)

suppressible

(Kim et al., 2012)

enhanceable

(Merigliano et al., 2017)

mei-41

enhanceable

tws

(Merigliano et al., 2017)

pan

enhanceable

tws

(Bajpai et al., 2004)

polo

enhanceable

tws

(Wang et al., 2011)

polo|gwl

enhanceable

tws

(Rangone et al., 2011)

Ret

enhanceable

tws

(Read et al., 2005)

sev|Ras85D

suppressible

tws

(Maixner et al., 1998)

sev|Ras85D

suppressible

tws|Delta

(Maixner et al., 1998)

sgg

enhanceable

tws

(Bajpai et al., 2004)

shg

suppressible

tws

(Greaves et al., 1999)

shg

enhanceable

tws

(Bajpai et al., 2004)

External Data

Subunit Structure (UniProtKB)

(UniProt, P36872 )

Linkouts

BioGRID - A database of protein and genetic interactions.

71024

DroID - A comprehensive database of gene and protein interactions.

FBgn0004889

MIST (genetic) - An integrated Molecular Interaction Database

47877

MIST (protein-protein) - An integrated Molecular Interaction Database

47877

Pathways

Signaling Pathways (FlyBase)

POSITIVE REGULATORS OF WNT-TCF SIGNALING PATHWAY
POSITIVE REGULATORS OF HEDGEHOG SIGNALING PATHWAY
POSITIVE REGULATORS OF TOLL-NF-kappaB SIGNALING PATHWAY

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.

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

P36872

Class of Gene

nuclear_gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

3-49

Cytogenetic map

85F13-85F14

Sequence location

3R:10,126,129..10,139,772 [-]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

85F13-85F14

Limits computationally determined from genome sequence between P{EP}Fmr1^EP3517&P{PZ}tws⁰²⁴¹⁴ and P{PZ}jumu⁰⁶⁴³⁹&P{lacW}l(3)j8B6^j8B6

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

85F12-85F16

(Mayer-Jaekel, 1993.5.7)

85F12-85F13

(determined by in situ hybridisation)

(Spradling et al., 1999, Perrimon et al., 1996)

85F11-85F16

(determined by in situ hybridisation)

(Szeged Stock Center, 1998-, Deak et al., 1997)

85F-85F

(determined by in situ hybridisation)

(Omel'yanchuk et al., 1997, Mayer-Jaekel et al., 1993, Uemura et al., 1993)

85E12-85E13

85F12--13

(BDGP Project Members, 1994-1999)

Experimentally Determined Recombination Data

Location

3-49

(FlyBase, 2016)

3-46.6

(Comeron et al., 2012)

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (30)

Aberrations including deletions of this gene Aberrations including duplications of this gene

Bloomington

49788

w¹¹¹⁸; P{GMR34E04-GAL4}attP2

50033

w¹¹¹⁸; P{GMR39A05-GAL4}attP2

54616

w¹¹¹⁸; P{GMR34E04-lexA}attP40

69054

w¹¹¹⁸; P{R34E04-GAL4.DBD}attP2

71342

w¹¹¹⁸; P{VT039305-p65.AD}attP40

Korea

12167

P{VT039305-GAL4}attP2

12453

P{VT039308-GAL4}attP2

15803

P{VT039305-lexA::GADfl}attP40

6235R-1

6235R-2

v104167

v34340

w¹¹¹⁸; P{GD10742}v34340

More stocks available...

Genomic Clones (18)

Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

cDNA Clones (146)

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

DPiM_LD12394

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.

FBgn0004889

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0004889

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Bajpai et al., 2004, Mayer-Jaekel et al., 1994, Uemura et al., 1993)

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

Other Comments

tws is required for efficient recruitment of pericentriolar material.

(Dobbelaere et al., 2008)

tws is required for the stabilization of arm in response to wg signaling.

(Bajpai et al., 2004)

dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R⁺ cell morphology.

(Kiger et al., 2003)

Identification: Enhancer trap expression pattern survey for loci expressed in the ring gland.

(Harvie et al., 1998)

Identification: Enhancer trap screen designed to discover genes involved in the cellular aspects of defense mechanisms, as well as in melanotic tumor formation processes linked to blood cell disregulation.

(Braun et al., 1997)

tws is involved in the control of cell division.

(Omel'yanchuk et al., 1997)

The autosomal "FLP-DFS" technique (using the P{ovoD1-18} P{FRT(w^hs)} P{hsFLP} chromosomes) has been used to identify the specific maternal effect phenotype for the zygotic lethal mutation.

(Perrimon et al., 1996)

Mutant alleles are pupal lethal and cause pattern duplication in imaginal discs.

(BDGP Project Members, 1994-1999)

The modulation of the phosphatase activity by the B subunit of type 2A protein phosphatase encoded by tws is crucial for specification of neural cell identity.

(Shiomi et al., 1994)

Mutations cause abnormalities in mitosis, both in the early embryo and in the larval brain.

(Gomes et al., 1993)

This locus encodes protein phosphatase 2A regulatory subunit. Mutants display intact lagging chromatids that have undergone separation from their sisters, but that remain at the position formerly occupied by the metaphase plate, as well as anaphase figures that show bridging chromatin having two centromeric regions.

(Mayer-Jaekel et al., 1993)

tws mutants display a mirror image duplication of parts of the imaginal wing disc suggesting that regulation of protein phosphatase activity may play a critical role in the specification of imaginal disc patterns.

(Uemura et al., 1993)

The tws gene product is a regulatory subunit of protein phosphatase 2 (PP2A), one of the major classes of serine/threonine phosphatases.

(Uemura et al., 1993)

tws is an essential gene required during embryogenesis and late larval development.

(Gomes et al., 1992)

Mutant alleles of tws are pupal lethal that display abnormal pattern formation in imaginal discs. tws gene product is highly homologous to the mammalian B subunits of protein phosphatase 2A.

(Uemura et al., 1992)

Relationship to Other Genes

Source for database merge of

Source for merge of: tws CG6235

(Bayraktaroglu, 2000.4.13)

Source for merge of: tws l(3)S061805

(Deak et al., 2003)

Source for merge of: l(3)S061805 l(3)S022361 l(3)S022440 l(3)S023013

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S035505b l(3)S043008b

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S075515b l(3)S022205b l(3)S023141a

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S134601a l(3)S146606 l(3)S091905

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S075902b l(3)S101413b l(3)S110008a

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S024834a l(3)S029701a l(3)S032708c

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S067915 l(3)S075110 l(3)S105605

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S060804 l(3)S061915 l(3)S110815

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S048013 l(3)S048507 l(3)S111515

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S042630 l(3)S043029 l(3)S119908

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S029403 l(3)S031006 l(3)S132907

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S026326 l(3)S027127 l(3)S141309

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S023206 l(3)S024838 l(3)S025806

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S060203 l(3)S067109b l(3)S069206a

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S076415 l(3)S080409 l(3)S088513

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S063110 l(3)S066017 l(3)S066813

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S049902 l(3)S052810 l(3)S053011

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S045519 l(3)S046918 l(3)S048006

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S032303 l(3)S033903 l(3)S042629

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S027313 l(3)S028707 l(3)S029110

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S025832 l(3)S025913 l(3)S026226

(Szeged Stock Center, 1998-)

Source for merge of: l(3)S061805 l(3)S023309 l(3)S023938 l(3)S024455

(Szeged Stock Center, 1998-)

Additional comments

Nomenclature History

Source for database identify of

Nomenclature comments

Etymology

Synonyms and Secondary IDs (104)

Reported As

Symbol Synonym

1466/06

(Deak et al., 1997)

2414

(Harvie et al., 1998)

B/PR55

(Janssens and Goris, 2001)

CG6235

(Çiçek et al., 2016, Hosono et al., 2015, White-Grindley et al., 2014, Ni et al., 2011.7.19, Ni et al., 2010.12.1, Jia et al., 2009, Keleman et al., 2009.8.5, Perkins et al., 2009, Wang et al., 2009, Dobbelaere et al., 2008, Hughes et al., 2008, Derré et al., 2007, Nybakken et al., 2005, Stempfl et al., 2002, Coulier et al., 2000)

DPR55

(Mayer-Jaekel and Hemmings, 1995)

PP2A

(Yeh et al., 2010, Bose et al., 2008, Sehgal, 2008, Su et al., 2008, Fang et al., 2007, Nam et al., 2007, Nam et al., 2007, Kyriacou and Odell, 2004, Bennett et al., 2003, Nowak et al., 2003, Bennett et al., 1999)

PP2A-B55

(Pechkovsky et al., 2013)

PP2A^B

(Kotadia et al., 2008)

PP2a

(Yu and Hardin, 2006)

PR55

(Wang et al., 2009, Millward et al., 1999, Mayer-Jaekel and Hemmings, 1994, Mayer-Jaekel et al., 1994, Mayer-Jaekel et al., 1993)

Pp2A-85F

TWS

(Avilés-Pagán et al., 2021)

Twins

(Brownlee and Rogers, 2013)

Tws

(Emond-Fraser et al., 2023, Neal et al., 2020, Zhao et al., 2017, Fischer et al., 2016, Brownlee et al., 2011, Rangone et al., 2011, Yeh et al., 2010, Jia et al., 2009, Krahn et al., 2009)

aar

(Pechkovsky et al., 2013, Lebedeva et al., 2003, Lebedeva et al., 2003, Lebedeva et al., 2000, Herrmann et al., 1999, Chubykin and Omelyanchuk, 1997, Chubykin et al., 1997, Omel'yanchuk et al., 1997, Craig and Brink, 1996, Mayer-Jaekel and Hemmings, 1994, Mayer-Jaekel et al., 1994, Williams and Goldberg, 1994, Gomes et al., 1993, Mayer-Jaekel et al., 1993, Mayer-Jaekel et al., 1993, Gomes et al., 1992)

dPP2A

(Merrow, 2006)

dPP2A-B55

(Pechkovsky et al., 2013)

dPP2A-Bα

(Bielinski and Mumby, 2007)

dPR55

(Li et al., 2002)

l(3)01436

l(3)02414

(Braun et al., 1997, Perrimon et al., 1996)

l(3)S022205b

l(3)S022361

l(3)S022440

l(3)S023013

l(3)S023141a

l(3)S023206

l(3)S023309

l(3)S023938

l(3)S024455

l(3)S024834a

l(3)S024838

l(3)S025806

l(3)S025832

l(3)S025913

l(3)S026226

l(3)S026326

l(3)S027127

l(3)S027313

l(3)S028707

l(3)S029110

l(3)S029403

l(3)S029701a

l(3)S031006

l(3)S032303

l(3)S032708c

l(3)S033903

l(3)S035505b

l(3)S042629

l(3)S042630

l(3)S043008b

l(3)S043029

l(3)S045519

l(3)S046918

l(3)S048006

l(3)S048013

l(3)S048507

l(3)S049902

(Read et al., 2005)

l(3)S052810

l(3)S053011

l(3)S060203

l(3)S060804

l(3)S061805

(Szeged Stock Center, 1998-)

l(3)S061915

l(3)S063110

l(3)S066017

l(3)S066813

l(3)S067109b

l(3)S067915

l(3)S069206a

l(3)S075110

l(3)S075515b

l(3)S075902b

l(3)S076415

l(3)S080409

l(3)S088513

l(3)S091905

l(3)S101413b

l(3)S105605

l(3)S110008a

l(3)S110815

l(3)S111515

l(3)S119908

l(3)S132907

l(3)S134601a

l(3)S141309

l(3)S146606

l(3)j11C8

l(3)s1801

(Jan and Jan, 1994)

tws

(Xu et al., 2025, Collins et al., 2024, Rangone et al., 2024, Barbaste et al., 2023, Praschberger et al., 2023, Chang et al., 2022, Miao et al., 2022, Perlegos et al., 2022, Barbosa et al., 2021, Järvelä-Stölting et al., 2021, Kanoh et al., 2021, Larouche et al., 2021, Lindsey et al., 2021, Ostalé et al., 2021, Pang et al., 2021, Garrido et al., 2020, Huang et al., 2020, Rui et al., 2020, Wolterhoff et al., 2020, Archambault, 2019.12.8, Kim et al., 2019, Rosen et al., 2019, Lee et al., 2018, Mehsen et al., 2018, Ashton-Beaucage and Therrien, 2017, Mendoza-Viveros et al., 2017, Merigliano et al., 2017, Clandinin and Owens, 2016-, Gene Disruption Project members, 2016-, Schwartz et al., 2016, Wang et al., 2016, Zhang et al., 2016, Garlapow et al., 2015, Gene Disruption Project members, 2015-, Liu and Secombe, 2015, Nagarkar-Jaiswal et al., 2015, Ashwal-Fluss et al., 2014, Faisal et al., 2014, Sopko et al., 2014, White-Grindley et al., 2014, Williams et al., 2014, Kwon et al., 2013, Pechkovsky et al., 2013, Bánréti et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Kim et al., 2012, Abruzzi et al., 2011, Friedman et al., 2011, Korcsmáros et al., 2011, Su et al., 2011, Wang et al., 2011, Wang et al., 2011, Yeh et al., 2010, Krahn et al., 2009, Moazzen et al., 2009, Wang et al., 2009, Batut et al., 2008, Cai et al., 2008, Christensen et al., 2008.6.11, Cook et al., 2008.9.29, Dobbelaere et al., 2008, Kotadia et al., 2008, Schuldiner et al., 2008, Yang et al., 2008, Bielinski and Mumby, 2007, Chen et al., 2007, Choi et al., 2007, Derré et al., 2007, Nybakken et al., 2005, Stanyon et al., 2004, Nowak et al., 2003)

tws/aar

(Deak et al., 2003)

v158

(Lebedeva et al., 2000)

Name Synonyms

PP2A regulatory subunit

(Jia et al., 2009)

PP2A regulatory subunit Twins

(Nie et al., 2009)

PP2A-B

(Fischer et al., 2016)

Protein phosphatase 2A at 85F

Twins

(Scholz, 2025, Choi et al., 2007, Li et al., 2002)

abnormal anaphase resolution

(Pechkovsky et al., 2013)

phosphoprotein phosphatase 2A 55 kDa

(Mayer-Jaekel, 1996.5.14, Mayer-Jaekel, 1996.5.14, Mayer-Jaekel, 1996.5.17, Mayer-Jaekel, 1993.5.7)

protein phosphatase 2A

(Yeh et al., 2010, Bose et al., 2008, Nam et al., 2007)

protein phosphatase 2a

(Yu and Hardin, 2006)

protein phosphatase type 2A

(Nowak et al., 2003)

regulatory B subunit of type 2A protein phosphatase

(Shiomi et al., 1994)

Secondary FlyBase IDs

FBgn0004906
FBgn0010249
FBgn0010748
FBgn0010780
FBgn0011012
FBgn0011434
FBgn0028118
FBgn0037791
FBgn0060325
FBgn0060355
FBgn0060382
FBgn0060393
FBgn0060451
FBgn0060476
FBgn0060482
FBgn0060490
FBgn0060503
FBgn0060530
FBgn0060599
FBgn0060620
FBgn0060667
FBgn0060680
FBgn0060683
FBgn0060685
FBgn0060686
FBgn0060717
FBgn0060727
FBgn0060736
FBgn0060738
FBgn0060744
FBgn0060764
FBgn0060771
FBgn0060777
FBgn0060782
FBgn0060814
FBgn0060817
FBgn0060835
FBgn0060851
FBgn0060857
FBgn0060858
FBgn0060872
FBgn0060877
FBgn0060907
FBgn0060908
FBgn0060915
FBgn0060916
FBgn0060954
FBgn0060957
FBgn0060959
FBgn0060962
FBgn0060965
FBgn0060971
FBgn0060972
FBgn0060974
FBgn0060977
FBgn0060995
FBgn0060996
FBgn0061006
FBgn0061008
FBgn0061016
FBgn0061017
FBgn0061018
FBgn0061025
FBgn0061026
FBgn0061030
FBgn0061036
FBgn0061050
FBgn0061051
FBgn0061053
FBgn0061055
FBgn0061059
FBgn0061060
FBgn0061062

Datasets (0)

Study focus (0)

Experimental Role

Project

Project Type

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Result

Result Type

Title

External Crossreferences and Linkouts ( 109 )

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.

47877

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.

P36872

UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information

P36872

UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information

A0A0B4LGZ3

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.

P36872

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0004889

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

47877/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0004889

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0004889

FlyMine - An integrated database for Drosophila genomics

FBgn0004889

InterPro - A database of protein families, domains and functional sites

KEGG Genes - Molecular building blocks of life in the genomic space.

dme:Dmel_CG6235

MARRVEL_MODEL - MARRVEL (model organism gene)

47877

Linkouts

BioGRID - A database of protein and genetic interactions.

71024

Cell Signaling Technology - Commercial vendor for primary antibodies and antibody conjugates.

2290

Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones

FBgn0004889

DroID - A comprehensive database of gene and protein interactions.

FBgn0004889

DRSC - Results frm RNAi screens

FBgn0004889

Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0004889

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0004889

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG6235

Flygut - An atlas of the Drosophila adult midgut

FBgn0004889

iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)

TC012273

Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution

Interactive Fly

KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.

MIST (genetic) - An integrated Molecular Interaction Database

47877

MIST (protein-protein) - An integrated Molecular Interaction Database

47877

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

P36872

References (241)

The following external sites may use different assemblies or annotations than FlyBase.

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