FB2022_03, released June 9, 2022

Gene: Dmel\gro

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General Information
Symbol
Dmel\gro
Species
D. melanogaster
Name
groucho
Annotation Symbol
CG8384
Feature Type
protein_coding_gene
FlyBase ID
FBgn0001139
Gene Model Status
Current
Stock Availability
22 publicly available
Gene Summary
groucho (gro) encodes a global developmental co-repressor in conjunction with manifold DNA-binding repressor partner proteins, which tether it to target promoters. It functions downstream of key signaling pathways such as Wg/Wnt and Dpp/TGF-beta. Notably, phosphorylation of the product of gro in response to MAPK activation weakens its repressor capacity. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Also Known As

m9/m10, m9/10, m9, m10, E(spl)m9/m10

Key Links
Genomic Location
Cytogenetic map
96F10-96F10
Sequence location
Recombination map
3-89
RefSeq locus
NT_033777 REGION:26041736..26049911
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
GO Summary Ribbons
Gene Ontology (GO) Annotations (19 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
enables CRD domain binding
inferred from physical interaction with UniProtKB:Q9UJU2
(assigned by UniProt )
(Arce et al., 2009)
enables DNA-binding transcription factor binding
inferred from physical interaction with FLYBASE:retn; FB:FBgn0004795
(Valentine et al., 1998)
inferred from physical interaction with FLYBASE:hkb; FB:FBgn0261434
(Goldstein et al., 1999)
inferred from physical interaction with FLYBASE:cic; FB:FBgn0262582
(Jimenez et al., 2000)
inferred from genetic interaction with FLYBASE:E(spl)m7-HLH; FB:FBgn0002633
(Giagtzoglou et al., 2003)
enables HMG box domain binding
inferred from physical interaction with UniProtKB:Q9UJU2
(assigned by UniProt )
(Arce et al., 2009)
enables protein binding
inferred from physical interaction with UniProtKB:P91943
(assigned by UniProt )
(Levanon et al., 1998)
inferred from physical interaction with UniProtKB:Q01196-1
(assigned by UniProt )
(Levanon et al., 1998)
inferred from physical interaction with UniProtKB:Q01196-2
(assigned by UniProt )
(Levanon et al., 1998)
inferred from physical interaction with UniProtKB:Q13761
(assigned by UniProt )
(Levanon et al., 1998)
inferred from physical interaction with UniProtKB:P14003
(assigned by UniProt )
(Fisher et al., 1996)
inferred from physical interaction with FLYBASE:elB; FB:FBgn0004858
(Dorfman et al., 2002)
inferred from physical interaction with FLYBASE:kni; FB:FBgn0001320
(Payankaulam and Arnosti, 2009)
inferred from physical interaction with FLYBASE:h; FB:FBgn0001168
(Zhang and Levine, 1999)
enables transcription corepressor activity
inferred from genetic interaction with FLYBASE:E(spl)m7-HLH; FB:FBgn0002633
(Giagtzoglou et al., 2003)
inferred from genetic interaction with FLYBASE:pan; FB:FBgn0085432
(Cavallo et al., 1998)
inferred from direct assay
(assigned by UniProt )
(Arce et al., 2009)
inferred from physical interaction with FLYBASE:h; FB:FBgn0001168
(Mannervik et al., 1999)
enables transcription factor binding
inferred from mutant phenotype
(Ajuria et al., 2011)
inferred from physical interaction with FLYBASE:HGTX; FB:FBgn0040318
(Syu et al., 2009)
inferred from physical interaction with FLYBASE:h; FB:FBgn0001168
(Goldstein et al., 1999)
inferred from physical interaction with FLYBASE:E(spl)m7-HLH; FB:FBgn0002633
(Goldstein et al., 1999)
inferred from physical interaction with FLYBASE:dl; FB:FBgn0260632
(Valentine et al., 1998)
inferred from physical interaction with FLYBASE:so; FB:FBgn0003460
(Kenyon et al., 2005)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
enables transcription corepressor activity
inferred from biological aspect of ancestor with PANTHER:PTN000850324
(assigned by GO_Central )
(Gaudet et al., 2011)
Biological Process (10 terms)
Terms Based on Experimental Evidence (9 terms)
CV Term
Evidence
References
involved_in epidermal growth factor receptor signaling pathway
inferred from genetic interaction with FLYBASE:Egfr; FB:FBgn0003731
(Johnston et al., 2016, Zhang and Du, 2015, Cinnamon et al., 2008)
inferred from mutant phenotype
(Helman et al., 2011)
involved_in fibroblast growth factor receptor signaling pathway
inferred from expression pattern
(Helman et al., 2011)
involved_in negative regulation of canonical Wnt signaling pathway
inferred from direct assay
(assigned by UniProt )
(Arce et al., 2009)
inferred from genetic interaction with FLYBASE:pan; FB:FBgn0085432
(Cavallo et al., 1998)
involved_in negative regulation of Toll signaling pathway
inferred from genetic interaction with FLYBASE:Tl; FB:FBgn0262473
(Koltun et al., 2017)
involved_in negative regulation of transcription by RNA polymerase II
inferred from mutant phenotype
(Zhang and Du, 2015)
inferred from physical interaction with FLYBASE:h; FB:FBgn0001168
(Zhang and Levine, 1999)
inferred from genetic interaction with FLYBASE:h; FB:FBgn0001168
(Forés et al., 2015)
involved_in negative regulation of transcription, DNA-templated
inferred from direct assay
(assigned by UniProt )
(Arce et al., 2009)
inferred from genetic interaction with FLYBASE:Myc; FB:FBgn0262656
(Orian et al., 2007)
involved_in nervous system development
inferred from mutant phenotype
(Schrons et al., 1992)
involved_in regulation of transcription, DNA-templated
inferred from mutant phenotype
(Payankaulam and Arnosti, 2009)
inferred from genetic interaction with FLYBASE:kni; FB:FBgn0001320
(Payankaulam and Arnosti, 2009)
involved_in torso signaling pathway
inferred from mutant phenotype
(Helman et al., 2011)
inferred from genetic interaction with FLYBASE:tor; FB:FBgn0003733
(Cinnamon et al., 2008)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
involved_in negative regulation of canonical Wnt signaling pathway
inferred from biological aspect of ancestor with PANTHER:PTN000850324
(assigned by GO_Central )
(Gaudet et al., 2011)
involved_in sex determination, primary response to X:A ratio
traceable author statement
(Schutt and Nothiger, 2000)
Cellular Component (3 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
part_of protein-containing complex
inferred from physical interaction with FLYBASE:Myc; FB:FBgn0262656
(Orian et al., 2007)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000850324
(assigned by GO_Central )
(Gaudet et al., 2011)
part_of transcription regulator complex
inferred from biological aspect of ancestor with PANTHER:PTN000850324
(assigned by GO_Central )
(Gaudet et al., 2011)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
Belongs to the WD repeat Groucho/TLE family. (P16371)
Protein Signatures (InterPro)
Summaries
Gene Snapshot
groucho (gro) encodes a global developmental co-repressor in conjunction with manifold DNA-binding repressor partner proteins, which tether it to target promoters. It functions downstream of key signaling pathways such as Wg/Wnt and Dpp/TGF-beta. Notably, phosphorylation of the product of gro in response to MAPK activation weakens its repressor capacity. [Date last reviewed: 2019-03-07]
Pathway (FlyBase)
Negative Regulators of Toll-NF-κB 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. Negative regulators of Toll-NF-κB Signaling lead to the increased cytosolic retention of the NF-κB proteins dl or Dif and/or repression of target gene transcription. (Adapted from FBrf0091014 and FBrf0223077).
Torso Signaling Pathway Core Components -
The formation of Drosophila embryonic termini is controlled by the localized activation of Torso (tor) receptor tyrosine kinase. The Torso signaling pathway acts via the canonical Ras/Raf/MAP kinase cascade. (Adapted from FBrf0157176.)
FGFR Signaling Pathway Core Components -
Fibroblast Growth Factor Receptor (FGFR) signaling pathway is initiated by the binding of secreted FGFs - bnl or ths/pyr to receptor tyrosine kinases btl or htl, respectively, to initiate signaling primarily via the canonical Ras/Raf/MAP kinase (ERK) cascade. (Adapted from FBrf0221038).
EGFR Signaling Pathway Core Components -
The Epidermal Growth Factor Receptor (EGFR) signaling pathway is used multiple times during development (FBrf0190321). It is activated by the binding of a secreted ligand to the receptor tyrosine kinase Egfr and acts via the canonical Ras/Raf/MAP kinase (ERK) cascade. (Adapted from FBrf0190321 and FBrf0221727).
Negative Regulators of Wnt-TCF Signaling Pathway -
Negative regulators of Wnt-TCF (canonical Wnt) signaling down-regulate the pathway, resulting in the attenuation of transcriptional regulation mediated by β-catenin (arm).
Protein Function (UniProtKB)
Transcriptional corepressor that regulates transcription when recruited to specific target DNA by hairy-related bHLH proteins. Maternally required for neurogenesis; in the segregation of the neuroectoderm. Directly or indirectly interacts with Notch and Delta.
(UniProt, P16371)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
E(spl)2
Homozygotes have clumps of extra bristles above each eye which give impression of bushy eyebrows; also extra bristles on the humerus. Top of head tends to be malformed; ocelli often enlarged and run together. In selected stocks, penetrance approaches 100%, but is low in unselected stocks. Concluded to be an allele of E(spl) on the basis of the visible phenotype of heterozygotes with lethal presumed point mutations at E(spl); however, it does not cause neural hypertrophy and is wild type in combination with E(spl)1.
Summary (Interactive Fly)

transcription factor - WD40 domain - E(spl) complex - partners transcriptional repressors by functioning as a co-repressor

Gene Model and Products
Number of Transcripts
10
Number of Unique Polypeptides
6

Please see the JBrowse view of Dmel\gro 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 New Section
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry P16371)

If you don't see the viewer to the right, 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.

Comments on Gene Model

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

Gene model reviewed during 5.48

Annotated transcripts do not represent all possible combinations of alternative exons and/or alternative promoters.

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

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

GC splice donor site postulated

Gene model reviewed during 5.50

Sequence Ontology: Class of Gene
nuclear_gene
protein_coding_gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
gro-RA
FBtr0084963
NM_170254
2988
719
gro-RB
FBtr0084962
NM_170255
2969
719
gro-RC
FBtr0084965
NM_170256
2887
719
gro-RD
FBtr0084964
NM_079790
2908
719
gro-RE
FBtr0084966
NM_206575
3002
730
gro-RF
FBtr0302951
NM_001202364
2975
719
gro-RG
FBtr0302952
NM_001202365
2996
728
gro-RH
FBtr0305047
NM_001260380
3951
709
gro-RI
FBtr0337057
NM_001276052
3918
698
gro-RJ
FBtr0339900
NM_001300610
3975
717
Additional Transcript Data and Comments
Reported size (kB)

4.4, 3.3 (northern blot)

(Knust et al., 1987)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
gro-PA
FBpp0084337
78.9
719
7.13
A0A0B4JD02
NP_733133
AAF56557
gro-PB
FBpp0084336
78.9
719
7.13
A0A0B4JD02
NP_733134
AAF56556
gro-PC
FBpp0084339
78.9
719
7.13
A0A0B4JD02
NP_733135
AAN14068
gro-PD
FBpp0084338
78.9
719
7.13
A0A0B4JD02
NP_524514
AAN14069
gro-PE
FBpp0089115
80.2
730
7.17
P16371
NP_996298
AAS65218
gro-PF
FBpp0292077
78.9
719
7.13
A0A0B4JD02
NP_001189293
ADV37383
gro-PG
FBpp0292078
80.0
728
7.17
A0A0B4JDF1
NP_001189294
ADV37384
gro-PH
FBpp0293584
77.9
709
8.03
A0A0B4K6L6
NP_001247309
AFH06626
gro-PI
FBpp0307986
76.6
698
8.02
A0A0B4KH09
NP_001262981
AGB96361
gro-PJ
FBpp0308932
78.7
717
7.13
A0A0B4LHP1
NP_001287539
AHN57538
Polypeptides with Identical Sequences

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

719 aa isoforms: gro-PA, gro-PB, gro-PC, gro-PD, gro-PF
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

Forms a complex with the hairy/Enhancer of split/deadpan family of basic helix-loop-helix proteins in order to repress transcription. Its activity in regulating transcription depends on other proteins as it lacks a DNA-binding motif.

(UniProt, P16371)
Post Translational Modification

Ubiquitinated by XIAP/BIRC4.

(UniProt, P16371)
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\gro 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
Expression Summary Ribbons
Colored tiles in ribbon indicate that the Fly Cell Atlas project found the gene expressed in that cell type. Darker colors mean that more cells of that cell type express the gene:
 low
high 
Colorless tiles indicate that there is no scRNAseq data for the gene in that cell type.
Colored tiles in ribbon indicate that expression data (RNA and/or protein) has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
Colored tiles in the ribbon indicate the average RNA expression level of the gene at the indicated stages:
 low
high 
as determined by RNA-seq (RPKM) using whole organism samples modENCODE, Brown et al., 2014. For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
central nervous system
(Maier et al., 1993)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
presumptive embryonic/larval peripheral nervous system
(Maier et al., 1993)
embryonic stage 1 -- 3
organism | ubiquitous

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 1 -- 5
organism
(Maier et al., 1993)
embryonic stage 4 -- 6
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 7 -- 8
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 9 -- 10
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 11 -- 12
central brain primordium
(Fisher et al., 2012)
ventral nerve cord primordium
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic brain
(Fisher et al., 2012)
larval ventral nerve cord
(Fisher et al., 2012)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 11
organism | ubiquitous
(Knust et al., 1987)
embryonic stage 5 -- 11

Comment: reference states 2-6 hr AEL

(Knust et al., 1987)
adult stage | female
(Knust et al., 1987)
Additional Descriptive Data

The distribution of embryonic gro transcripts was compared in D. melanogaster and D. hydei. The patterns of embryonic gene activity were found to be nearly indistinguishable.

(Maier et al., 1993)

gro transcripts are expressed at high levels in 2-6 hr embryos, and in adult females. In situ hybridization shows that gro transcripts are ubiquitous in the embryo, and persist until late stage 11.

(Knust et al., 1987)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 9
embryonic neuroblast
(Orian et al., 2007)
adult stage | female
germarium
(Parrott et al., 2011)
female germline cell
(Parrott et al., 2011)
adult stage | female & oogenesis
prefollicle cell
(Johnston et al., 2016)
escort cell
(Johnston et al., 2016)
female germline cell
(Johnston et al., 2016)
Additional Descriptive Data

gro protein has strong expression in all follicle cells in the germarium, and more weakly in germ cells and escort cells.

(Johnston et al., 2016)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
part_of protein-containing complex
inferred from physical interaction with FLYBASE:Myc; FB:FBgn0262656
(Orian et al., 2007)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\gro in GBrowse 2
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
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 68 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 49 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of gro
Transgenic constructs containing regulatory region of gro
Aberrations (Deficiencies and Duplications) ( 36 )
Inferred from experimentation ( 36 )
Gene disrupted in
Df(3R)boss9
(Fischer-Vize et al., 1992)
Df(3R)BSC751
(Christensen et al., 2009.2.28)
Df(3R)E(spl)-r8.1
(Heitzler et al., 1996, Schrons et al., 1992)
Df(3R)E(spl)-r72.1
(Schrons et al., 1992)
Df(3R)E(spl)-rv27
(de Celis and Ruiz-Gomez, 1995, Tietze et al., 1993, Schrons et al., 1992)
Df(3R)ED6232
(Ryder and Roote, 2004.11.10)
Df(3R)Espl1
(Bodmer et al., 1993)
Df(3R)Espl3
(Crews, 2007.1.6, Jiang and Crews, 2006, Cook and Carpenter, 2002)
Df(3R)Espl4
(Heitzler et al., 1996, Schrons et al., 1992)
Df(3R)Espl9
(de Celis and Ruiz-Gomez, 1995, Fischer-Vize et al., 1992)
Df(3R)Espl10
(Fischer-Vize et al., 1992)
Df(3R)Espl22
(Ligoxygakis et al., 1998, Price et al., 1997, de Celis and Ruiz-Gomez, 1995, Fischer-Vize et al., 1992)
Df(3R)Exel6204
(Crews, 2007.1.6, Ryder, 2004.4.26)
Df(3R)grob32.2
(de Celis et al., 1997, Heitzler et al., 1996)
Df(3R)gror32.1
(Schrons et al., 1992)
Df(3R)grory78R-r171.1
(Schrons et al., 1992, Tietze et al., 1992)
Df(3R)P709
(Fischer-Vize et al., 1992, Delidakis et al., 1991)
Df(3R)R20
(Wurmbach and Preiss, 2014)
Df(3R)R70
(Wurmbach and Preiss, 2014)
Df(3R)R75
(Wurmbach and Preiss, 2014)
Df(3R)ro80b
(Cook and Carpenter, 2002)
Df(3R)Tl-I
(Cook and Carpenter, 2002)
Df(3R)Tl-P
(Cook and Carpenter, 2002)
Df(3R)X13
(Wurmbach and Preiss, 2014)
T(2,3)R72
(Wurmbach and Preiss, 2014)
T(2;3)boss3
(Fischer-Vize et al., 1992)
Gene partially disrupted in
Df(3R)grob32.2
(Ligoxygakis et al., 1998, de Celis et al., 1996, Schrons et al., 1992)
Gene not disrupted in
Df(3R)boss14
(Delidakis et al., 1991)
Df(3R)boss15
(Fischer-Vize et al., 1992, Delidakis et al., 1991)
Df(3R)boss16
(Fischer-Vize et al., 1992)
Df(3R)E(spl)δ-6
(Bardin et al., 2010, Chanet et al., 2009)
Df(3R)ED6235
(Ryder and Roote, 2004.11.10)
Df(3R)Espl-X10
(Wurmbach and Preiss, 2014)
Df(3R)ME61
(Cook and Carpenter, 2002)
Df(3R)Tl-X
(Cook and Carpenter, 2002)
Gene duplicated in
Dp(3;2)E(spl)δ-8Δα46
(Chanet et al., 2009)
Dp(3;2)E(spl)δ-8
(Chanet et al., 2009)
Inferred from location ( 0 )
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\GAL429BD
lethal, with Scer\GAL4c355
lethal, with Scer\GAL4dpp.blk1
lethal, with Scer\GAL4hs.2sev
lethal (with Df(3R)R20)
lethal (with Df(3R)R27)
lethal (with Df(3R)R70)
lethal (with Df(3R)R75)
lethal (with Df(3R)X13)
lethal (with T(2,3)R72)
lethal | dominant | maternal effect
lethal | embryonic stage | recessive
lethal | maternal effect, with Scer\GAL4VP16.nos.UTR
lethal | neurogenic phenotype
lethal | recessive
lethal - all die before end of P-stage, with Scer\GAL4bi-omb-Gal4
lethal - all die before end of pupal stage | recessive
lethal - all die during embryonic stage, with Scer\GAL4VP16.mat.αTub67C
lethal - all die during embryonic stage | maternal effect, with Scer\GAL4VP16.nos.UTR
lethal - all die during embryonic stage | neurogenic phenotype | recessive
lethal - all die during embryonic stage | recessive
partially lethal - majority die, with Scer\GAL4pnr-MD237
partially lethal - majority live
viable
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4tin.CΔ4
viable (with Df(3R)Espl-X10)
viable | recessive
Sterility
Allele
female semi-sterile
male semi-sterile
Other Phenotypes
Allele
abnormal neuroanatomy | embryonic stage, with Scer\GAL4VP16.nos.UTR
embryonic/larval segmentation phenotype | maternal effect, with Scer\GAL4VP16.nos.UTR
increased cell number | oogenesis | oogenesis, with Scer\GAL4109-30
some die during embryonic stage, with Scer\GAL4VP16.mat.αTub67C
some die during embryonic stage | maternal effect, with Scer\GAL4VP16.nos.UTR
some die during embryonic stage | recessive
some die during pharate adult stage, with Scer\GAL4bi-omb-Gal4
some die during pupal stage, with Scer\GAL4pnr-MD237
some die during pupal stage | recessive
visible
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4ey.PH
visible, with Scer\GAL4ey.PU
visible, with Scer\GAL4GMR.PU
visible, with Scer\GAL4pnr-MD237
visible, with Scer\GAL4sd-SG29.1
visible, with Scer\GAL4Ser.PGF
visible (with gro1)
visible (with groC105)
visible | dominant
visible | recessive
visible | recessive | somatic clone
visible | somatic clone
wild-type
Phenotype manifest in
Allele
adult head & microchaeta
anterior postalar bristle
basal stalk, with Scer\GAL4109-30
chaeta | ectopic | somatic clone
chaeta | increased number, with Scer\GAL4pnr-MD237
dendrite
denticle belt
dorsal-ventral compartment boundary of the wing blade
embryonic/first instar larval cuticle
embryonic/first instar larval cuticle, with Scer\GAL4VP16.mat.αTub67C
embryonic/first instar larval cuticle | germline clone | maternal effect
embryonic/first instar larval cuticle | maternal effect
embryonic epidermis
embryonic gnathal segment
eye
eye, with Scer\GAL4ey.PH
eye, with Scer\GAL4ey.PU
eye, with Scer\GAL4GMR.PU
eye-antennal disc | late third instar larval stage, with Scer\GAL4ey.PH
eye photoreceptor cell & eye-antennal disc, with Scer\GAL4ey.PH
filzkorper | maternal effect, with Scer\GAL4VP16.nos.UTR
follicle cell, with Scer\GAL4109-30
follicle stem cell | somatic clone, with Scer\GAL4Tub.PU
frontal bristle | increased number (with gro1)
frontal bristle | increased number (with groC105)
germarium, with Scer\GAL4109-30
germarium | somatic clone, with Scer\GAL4Tub.PU
head capsule, with Scer\GAL4ey.PH
intermediate ventral neurectoderm, with Scer\GAL4VP16.nos.UTR
larval head | maternal effect, with Scer\GAL4VP16.nos.UTR
leg, with Scer\GAL4bi-omb-Gal4
macrochaeta
macrochaeta & abdominal tergite
macrochaeta & head
macrochaeta & scutum
macrochaeta & thorax
mesothoracic tergum, with Scer\GAL4pnr-MD237
mesothoracic tergum & macrochaeta, with Scer\GAL4sca-C253
mesothoracic tergum | somatic clone
microchaeta & abdominal tergite
microchaeta & head
microchaeta & scutum
microchaeta & thorax
microchaeta | ectopic | somatic clone
ocellus
ommatidium
photoreceptor neuron | somatic clone
scutellum | somatic clone
sense organ & wing
sensory mother cell, with Scer\GAL4sca-C253
stalk follicle cell, with Scer\GAL4109-30
ventral nerve cord primordium, with Scer\GAL4rho.PU
ventral nerve cord primordium | maternal effect
wing, with Scer\GAL4Bx-MS1096
wing, with Scer\GAL4sd-SG29.1
wing, with Scer\GAL4Ser.PGF
wing blade anterior compartment
wing margin, with Scer\GAL4sd-SG29.1
wing vein, with Scer\GAL4Bx-MS1096
wing vein, with Scer\GAL4sd-SG29.1
wing vein L1
wing vein L2
wing vein L3
wing vein L4, with Scer\GAL4Ser.PGF
wing vein L5
wing vein L5, with Scer\GAL4Ser.PGF
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (7)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Hsap\TLE3
12 of 15
Yes
Yes
Hsap\TLE4
12 of 15
Yes
Yes
Hsap\TLE1
11 of 15
No
Yes
3  
Hsap\TLE2
11 of 15
No
Yes
Hsap\TLE5
4 of 15
No
Yes
1  
Hsap\TLE6
4 of 15
No
Yes
Hsap\TLE7
2 of 15
No
Yes
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (7)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Mmus\Tle3
12 of 15
Yes
Yes
Mmus\Tle4
12 of 15
Yes
Yes
Mmus\Tle2
11 of 15
No
Yes
Mmus\Tle1
10 of 15
No
Yes
Mmus\Tle5
4 of 15
No
Yes
Mmus\Tle6
4 of 15
No
Yes
Mmus\Tle7
3 of 15
No
Yes
Rattus norvegicus (Norway rat) (7)
Rnor\Tle3
10 of 13
Yes
Yes
Rnor\Tle1
9 of 13
No
Yes
Rnor\Tle4
8 of 13
No
Yes
Rnor\Tle2
6 of 13
No
Yes
Rnor\Tle6
4 of 13
No
Yes
Rnor\Tle5
2 of 13
No
Yes
Rnor\Tle7
2 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (4)
Xtro\tle1
8 of 12
Yes
Yes
Xtro\tle2
7 of 12
No
Yes
Xtro\tle4
7 of 12
No
Yes
Xtro\aes
2 of 12
No
Yes
Danio rerio (Zebrafish) (7)
Drer\tle3b
13 of 15
Yes
Yes
Drer\tle3a
12 of 15
No
Yes
Drer\tle2a
11 of 15
No
Yes
Drer\tle2b
9 of 15
No
Yes
Drer\chico
3 of 15
No
Yes
Drer\tle5
3 of 15
No
Yes
Drer\tle2c
2 of 15
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (4)
Cele\unc-37
10 of 15
Yes
Yes
Cele\eya-1
1 of 15
No
No
Cele\wdr-5.3
1 of 15
No
No
Cele\wdr-48
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (1)
Atha\SCD1
1 of 9
Yes
Yes
Saccharomyces cerevisiae (Brewer's yeast) (1)
Scer\TUP1
1 of 15
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Other Organism Orthologs (via OrthoDB)
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (1)
Gene Symbol
Score
Source
Compara
eggNOG
Homologene
Inparanoid
Isobase
OrthoDB
OrthoFinder
orthoMCL
Panther
RoundUp
Alignment
CG5466
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 1 )
    Allele
    Disease
    Interaction
    References
    gro1
    ameliorates  Schimke immuno-osseous dysplasia
    modeled by Marcal1UAS.cBa
    (Morimoto et al., 2016)
    Disease Associations of Human Orthologs (via DIOPT v8.0 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?
    TLE3; TLE family member 3, transcriptional corepressor
    12 of 15
    600190
        TLE4; TLE family member 4, transcriptional corepressor
        12 of 15
        605132
            TLE1; TLE family member 1, transcriptional corepressor
            11 of 15
            600189
                 
                TLE2; TLE family member 2, transcriptional corepressor
                11 of 15
                601041
                    TLE5; TLE family member 5, transcriptional modulator
                    4 of 15
                    600188
                         
                        TLE6; TLE family member 6, subcortical maternal complex member
                        4 of 15
                        612399
                          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
                          esyN Network Diagram
                          Show neighbor-neighbor interactions:
                          Select Layout:
                          Legend:
                          Protein
                          RNA
                          Selected Interactor(s)
                          Interactions Browser
                          View in Interactions Browser
                          Please see the Physical Interaction reports below for full details
                          protein-protein
                          Physical Interaction
                          Assay
                          References
                          gro
                          molecular sieving, molecular weight estimation by staining, blue native page, autoradiography, pull down, anti tag coimmunoprecipitation, western blot, affinity chromatography technology, cosedimentation, molecular weight
                          (Turki-Judeh and Courey, 2012, Kuo et al., 2011, Jennings et al., 2008, Mieszczanek et al., 2008, Song et al., 2004)
                          gro - Ada2b
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - Alh
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - B-H1
                          pull down, autoradiography
                          (Goldstein et al., 2005)
                          gro - Boh1
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - Chi
                          anti tag coimmunoprecipitation, anti tag western blot
                          (Fiedler et al., 2015)
                          gro - CtBP
                          anti bait coimmunoprecipitation, western blot
                          (Nagel et al., 2005)
                          gro - D
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - Dlip2
                          two hybrid
                          (Bhaskar et al., 2000)
                          gro - Doc1
                          pull down, autoradiography, two hybrid
                          (Goldstein et al., 2005)
                          gro - E(spl)m3-HLH
                          two hybrid, pull down, autoradiography, inferred by curator
                          (Jennings et al., 2006, Poortinga et al., 1998, Alifragis et al., 1997, Paroush et al., 1994)
                          gro - E(spl)m4-BFM
                          two hybrid
                          (Poortinga et al., 1998)
                          gro - E(spl)m5-HLH
                          two hybrid, pull down, autoradiography, inferred by curator
                          (Jennings et al., 2006, Poortinga et al., 1998, Alifragis et al., 1997, Paroush et al., 1994)
                          gro - E(spl)m7-HLH
                          anti tag coimmunoprecipitation, western blot, pull down, autoradiography, two hybrid, inferred by curator
                          (Kiparaki et al., 2015, Zarifi et al., 2012, Bejarano et al., 2007, Jennings et al., 2006, Giagtzoglou et al., 2005, Goldstein et al., 1999, Levanon et al., 1998, Alifragis et al., 1997, Fisher et al., 1996, Paroush et al., 1994)
                          gro - E(spl)m8-HLH
                          two hybrid, anti tag coimmunoprecipitation, western blot, pull down, autoradiography, inferred by curator
                          (Kiparaki et al., 2015, Eastwood et al., 2011, Kahali et al., 2010, Jennings et al., 2006, Karandikar et al., 2004, Poortinga et al., 1998, Alifragis et al., 1997, Paroush et al., 1994)
                          gro - E(spl)mβ-HLH
                          two hybrid, pull down, autoradiography, inferred by curator
                          (Jennings et al., 2006, Poortinga et al., 1998, Alifragis et al., 1997)
                          gro - E(spl)mγ-HLH
                          pull down, autoradiography, inferred by curator, two hybrid
                          (Jennings et al., 2006, Poortinga et al., 1998, Alifragis et al., 1997)
                          gro - E(spl)mδ-HLH
                          two hybrid array, pull down, autoradiography, inferred by curator, two hybrid
                          (Shokri et al., 2019, Jennings et al., 2006, Poortinga et al., 1998, Alifragis et al., 1997, Paroush et al., 1994)
                          gro - Eip78C
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - Ets65A
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - H
                          two hybrid, pull down, autoradiography, anti bait coimmunoprecipitation, western blot
                          (Nagel et al., 2020, Zehender et al., 2017, Nagel and Preiss, 2014, Nagel et al., 2005, Barolo et al., 2002)
                          gro - HDAC1
                          pull down, western blot, autoradiography, anti tag coimmunoprecipitation, anti bait coimmunoprecipitation, peptide massfingerprinting
                          (Choi et al., 2005, Chen et al., 1999)
                          gro - HGTX
                          anti tag coimmunoprecipitation, anti tag western blot
                          (Syu et al., 2009)
                          gro - Hesr
                          pull down, autoradiography, inferred by curator
                          (Jennings et al., 2006)
                          gro - Him
                          pull down, autoradiography, inferred by curator
                          (Liotta et al., 2007, Jennings et al., 2006)
                          gro - Hipk
                          protein kinase assay, autoradiography, bimolecular fluorescence complementation, fluorescence microscopy, enzymatic study
                          (Steinmetz et al., 2018, Dewald et al., 2014, Lee et al., 2009, Choi et al., 2005)
                          gro - His1
                          anti tag coimmunoprecipitation, peptide massfingerprinting
                          (Chen et al., 1999)
                          gro - His3
                          pull down, autoradiography
                          (Turki-Judeh and Courey, 2012)
                          gro - His4
                          pull down, autoradiography
                          (Turki-Judeh and Courey, 2012)
                          gro - Hr96
                          pull down, autoradiography
                          (Goldstein et al., 2005)
                          gro - Irbp18
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - Kr-h1
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - Myc
                          pull down, autoradiography, anti bait coimmunoprecipitation, western blot, anti tag western blot
                          (Orian et al., 2007)
                          gro - Optix
                          two hybrid, anti tag coimmunoprecipitation, anti tag western blot
                          (Anderson et al., 2012, Kenyon et al., 2005)
                          gro - Pep
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - Sidpn
                          pull down, autoradiography, inferred by curator
                          (Jennings et al., 2006)
                          gro - Su(var)2-10
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - Ubx
                          bimolecular fluorescence complementation, fluorescence microscopy
                          (Bischof et al., 2018)
                          gro - asf1
                          anti bait coimmunoprecipitation, western blot
                          (Goodfellow et al., 2007)
                          gro - bap
                          pull down, autoradiography
                          (Goldstein et al., 2005)
                          gro - bowl
                          two hybrid, two hybrid array
                          (Shokri et al., 2019, Goldstein et al., 2005)
                          gro - brk
                          pull down, autoradiography, two hybrid
                          (Turki-Judeh and Courey, 2012, Ziv et al., 2009, Song et al., 2004, Hasson et al., 2001)
                          gro - cic
                          pull down, autoradiography
                          (Jimenez et al., 2000)
                          gro - dl
                          anti tag coimmunoprecipitation, western blot, pull down, autoradiography
                          (Dubnicoff et al., 1997)
                          gro - dpn
                          two hybrid, pull down, autoradiography, inferred by curator
                          (Jennings et al., 2006, Poortinga et al., 1998, Paroush et al., 1994)
                          gro - elB
                          pull down, autoradiography
                          (Dorfman et al., 2002)
                          gro - ems
                          pull down, autoradiography
                          (Goldstein et al., 2005)
                          gro - en
                          pull down, autoradiography, anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based, two hybrid
                          (Rhee et al., 2014, Hittinger and Carroll, 2008, Jennings et al., 2006, Goldstein et al., 2005, Kobayashi et al., 2001, Tolkunova et al., 1998, Jimenez et al., 1997)
                          gro - eve
                          pull down, autoradiography, two hybrid
                          (Braid et al., 2010, Kobayashi et al., 2001)
                          gro - ey
                          pull down, autoradiography, anti tag coimmunoprecipitation, anti tag western blot
                          (Choi et al., 2005)
                          gro - eya
                          pull down, autoradiography
                          (Goldstein et al., 2005)
                          gro - ftz-f1
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - h
                          pull down, autoradiography, two hybrid, anti tag western blot, western blot
                          (Abed et al., 2011, Payankaulam and Arnosti, 2009, Bejarano et al., 2007, Orian et al., 2007, Jennings et al., 2006, Goldstein et al., 2005, Barolo et al., 2002, Hasson et al., 2001, Jimenez et al., 2000, Goldstein et al., 1999, Zhang and Levine, 1999, Poortinga et al., 1998, Alifragis et al., 1997, Jimenez et al., 1997, Fisher et al., 1996, Paroush et al., 1994)
                          gro - hkb
                          two hybrid, pull down, autoradiography
                          (Bejarano et al., 2007, Goldstein et al., 1999)
                          gro - ind
                          anti tag coimmunoprecipitation, anti tag western blot
                          (Von Ohlen and Moses, 2009, Von Ohlen et al., 2007)
                          gro - kni
                          pull down, autoradiography, western blot, anti tag coimmunoprecipitation
                          (Payankaulam and Arnosti, 2009)
                          gro - mid
                          anti bait coimmunoprecipitation, western blot, anti tag coimmunoprecipitation, pull down, autoradiography
                          (Formaz-Preston et al., 2012)
                          gro - odd
                          two hybrid, pull down, autoradiography
                          (Goldstein et al., 2005)
                          gro - pan
                          pull down, autoradiography
                          (Mieszczanek et al., 2008, Levanon et al., 1998)
                          gro - row
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - run
                          pull down, western blot, two hybrid
                          (Walrad et al., 2011, Alifragis et al., 1997, Aronson et al., 1997)
                          gro - salm
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - sbb
                          anti bait coimmunoprecipitation, anti tag western blot
                          (Bejarano et al., 2007)
                          gro - sc
                          anti tag coimmunoprecipitation, anti tag western blot
                          (Kiparaki et al., 2015)
                          gro - shn
                          anti tag coimmunoprecipitation, anti tag western blot
                          (Cai and Laughon, 2009)
                          gro - slp1
                          two hybrid, pull down, autoradiography
                          (Goldstein et al., 2005, Kobayashi et al., 2001)
                          gro - so
                          two hybrid
                          (Kenyon et al., 2005, Kenyon et al., 2005)
                          gro - tio
                          two hybrid array
                          (Shokri et al., 2019)
                          gro - vnd
                          anti tag coimmunoprecipitation, anti tag western blot, anti bait coimmunoprecipitation, western blot, pull down, autoradiography
                          (Zhang et al., 2008, Uhler et al., 2007, Goldstein et al., 2005)
                          RNA-protein
                          Physical Interaction
                          Assay
                          References
                          gro - snRNP-U1-C
                          coimmunoprecipitation, quantitative reverse transcription pcr
                          (Kwong et al., 2015)
                          Summary of Genetic Interactions
                          esyN Network Diagram
                          esyN Network Key:
                          Suppression
                          Enhancement
                          View in Interactions Browser
                          Please look at the allele data for full details of the genetic interactions
                          Starting gene(s)
                          Interaction type
                          Interacting gene(s)
                          Reference
                          gro
                          suppressible
                          btl::Egfr
                          (Hasson et al., 2005)
                          gro
                          enhanceable
                          CG12299
                          (Haussmann et al., 2008)
                          gro|h
                          suppressible
                          CtBP
                          (Phippen et al., 2000)
                          gro
                          suppressible
                          dgrn
                          (Abed et al., 2011)
                          gro
                          enhanceable
                          E(spl)mβ-HLH
                          (Hasson et al., 2005)
                          gro
                          enhanceable
                          Egfr
                          (Price et al., 1997)
                          gro
                          suppressible
                          Hipk
                          (Lee et al., 2009, Choi et al., 2005)
                          gro
                          suppressible
                          klu
                          (Kaspar et al., 2008)
                          gro
                          enhanceable
                          kni
                          (Payankaulam and Arnosti, 2009)
                          gro
                          suppressible
                          N
                          (Ramain et al., 2001)
                          gro
                          enhanceable
                          rl
                          (Price et al., 1997)
                          gro
                          enhanceable
                          wg
                          (Lawrence et al., 2000)
                          Starting gene(s)
                          Interaction type
                          Interacting gene(s)
                          Reference
                          al
                          enhanceable
                          gro
                          (Wehn and Campbell, 2006)
                          arm
                          suppressible
                          gro
                          (Cavallo et al., 1998)
                          btl::Egfr
                          suppressible
                          gro
                          (Johnston et al., 2016)
                          Chi
                          suppressible
                          gro
                          (van Tienen et al., 2017)
                          Egfr
                          enhanceable
                          gro
                          (Price et al., 1997)
                          eve
                          enhanceable
                          gro
                          (Kobayashi et al., 2001)
                          h
                          suppressible
                          gro
                          (Jennings et al., 2008)
                          H
                          enhanceable
                          gro
                          (Lee et al., 2009, Barolo et al., 2002, Johannes and Preiss, 2002)
                          h
                          enhanceable
                          gro
                          (Phippen et al., 2000)
                          H
                          suppressible
                          gro
                          (Lee et al., 2009, Johannes and Preiss, 2002)
                          Him
                          suppressible
                          gro
                          (Liotta et al., 2007)
                          kni
                          enhanceable
                          gro
                          (Payankaulam and Arnosti, 2009)
                          mid
                          enhanceable
                          gro
                          (Formaz-Preston et al., 2012)
                          N
                          suppressible
                          gro
                          (Barolo et al., 2002)
                          pan
                          suppressible
                          gro
                          (Lawrence et al., 2000, Cavallo et al., 1998)
                          ro
                          suppressible
                          gro
                          (Chanut et al., 2000)
                          run
                          suppressible
                          gro
                          (Wheeler et al., 2002)
                          Snr1
                          enhanceable
                          gro
                          (Marenda et al., 2004)
                          Su(H)
                          suppressible
                          gro
                          (Price et al., 1997)
                          Su(H)
                          enhanceable
                          gro
                          (Barolo et al., 2002)
                          External Data
                          Subunit Structure (UniProtKB)
                          Forms a complex with the hairy/Enhancer of split/deadpan family of basic helix-loop-helix proteins in order to repress transcription. Its activity in regulating transcription depends on other proteins as it lacks a DNA-binding motif.
                          (UniProt, P16371 )
                          Linkouts
                          BioGRID - A database of protein and genetic interactions.
                          DroID - A comprehensive database of gene and protein interactions.
                          MIST (genetic) - An integrated Molecular Interaction Database
                          MIST (protein-protein) - An integrated Molecular Interaction Database
                          Pathways
                          Signaling Pathways (FlyBase)
                          Negative Regulators of Toll-NF-κB 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. Negative regulators of Toll-NF-κB Signaling lead to the increased cytosolic retention of the NF-κB proteins dl or Dif and/or repression of target gene transcription. (Adapted from FBrf0091014 and FBrf0223077).
                          Torso Signaling Pathway Core Components -
                          The formation of Drosophila embryonic termini is controlled by the localized activation of Torso (tor) receptor tyrosine kinase. The Torso signaling pathway acts via the canonical Ras/Raf/MAP kinase cascade. (Adapted from FBrf0157176.)
                          FGFR Signaling Pathway Core Components -
                          Fibroblast Growth Factor Receptor (FGFR) signaling pathway is initiated by the binding of secreted FGFs - bnl or ths/pyr to receptor tyrosine kinases btl or htl, respectively, to initiate signaling primarily via the canonical Ras/Raf/MAP kinase (ERK) cascade. (Adapted from FBrf0221038).
                          EGFR Signaling Pathway Core Components -
                          The Epidermal Growth Factor Receptor (EGFR) signaling pathway is used multiple times during development (FBrf0190321). It is activated by the binding of a secreted ligand to the receptor tyrosine kinase Egfr and acts via the canonical Ras/Raf/MAP kinase (ERK) cascade. (Adapted from FBrf0190321 and FBrf0221727).
                          Negative Regulators of Wnt-TCF Signaling Pathway -
                          Negative regulators of Wnt-TCF (canonical Wnt) signaling down-regulate the pathway, resulting in the attenuation of transcriptional regulation mediated by β-catenin (arm).
                          Metabolic Pathways
                          External Data
                          Genomic Location and Detailed Mapping Data
                          Chromosome (arm)
                          3R
                          Recombination map
                          3-89
                          Cytogenetic map
                          96F10-96F10
                          Sequence location
                          FlyBase Computed Cytological Location
                          Cytogenetic map
                          Evidence for location
                          96F10-96F10
                          Limits computationally determined from genome sequence between P{PZ}l(3)rQ197rQ197 and P{lacW}scribj7B3
                          Experimentally Determined Cytological Location
                          Cytogenetic map
                          Notes
                          References
                          96F10-96F11
                          (Price et al., 1997)
                          96F11-96F14
                          (determined by in situ hybridisation)
                          (Hartley et al., 1988)
                          Experimentally Determined Recombination Data
                          Location

                          3-89

                          (FlyBase, 2016)

                          3-87.4

                          (Comeron et al., 2012)

                          3-87

                          (Price et al., 1997)

                          3-89.1

                          Left of (cM)
                          Right of (cM)
                          Notes
                          Stocks and Reagents
                          Stocks (22)
                          Genomic Clones (25)
                          cDNA Clones (145)
                           

                          Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see GBrowse for alignment of the cDNAs and ESTs to the gene model.

                          cDNA clones, fully sequenced
                          BDGP DGC clones
                          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
                          GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
                          Antibody Information
                          Laboratory Generated Antibodies
                           

                          polyclonal

                          (Chambers et al., 2017, Kwong et al., 2015, Cinnamon et al., 2008, Choi et al., 2005)
                          Commercially Available Antibodies
                           
                          Cell Signaling Technology - Commercial vendor for primary antibodies and antibody conjugates.
                          Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
                          Other Information
                          Relationship to Other Genes
                          Source for database identify of
                          Source for database merge of

                          Source for merge of: gro BEST:LD15161 BEST:GM01575

                          (Rana, 2004.7.8)

                          Source for merge of: gro BcDNA:LD33829

                          (FlyBase, 1996-)

                          Source for merge of: gro anon-WO0118547.385

                          (FlyBase, 1996-)
                          Additional comments

                          Source for merge of gro BcDNA:LD33829 was sequence comparison ( date:990717 ).

                          (FlyBase, 1996-)

                          Source for merge of gro anon-WO0118547.385 was sequence comparison ( date:051113 ).

                          (FlyBase, 1996-)
                          Other Comments

                          DNA-protein interactions: genome-wide binding profile assayed for gro protein in Kc167 cells; see Chromatin_types_NKI collection report. Individual protein-binding experiments listed under "Samples" at GEO_GSE22069 (http://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE22069).

                          (Filion et al., 2010)

                          dsRNA made from templates generated with primers directed against this gene has been transfected into Kc cells.

                          (Fang et al., 2006)

                          dsRNA has been made from templates generated with primers directed against this gene. RNAi of gro results in dorsal overextension of primary dendrites and a reduction in lateral branching.

                          (Parrish et al., 2006)

                          gro forms a junction between signalling pathways, enabling Egfr signalling to antagonize transcriptional output by N.

                          (Hasson et al., 2005)

                          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)

                          RNAi screen using dsRNA made from templates generated with primers directed against this gene causes a phenotype when assayed in Kc167 and S2R+ cells: increased or polarized (uneven) accumulation of F-actin.

                          (Kiger et al., 2003)

                          gro mediates a ci-independant mechanism of hh repression in the anterior wing pouch.

                          (Apidianakis et al., 2001)

                          Area matching Drosophila EST AA540693. This EST forms a 1450bp contig with ESTs AA441321, AA440080 and AA392794.

                          (Davies, 2001.3.15)

                          The brk co-repressors gro and CtBP are alternately used for repressing some dpp-responsive genes, whereas for repressing other, distinct, target genes they are not required.

                          (Hasson et al., 2001)

                          gro and H control different aspects of ato's complex expression pattern.

                          (Chanut et al., 2000)

                          gro and H both act to restrict neuronal cell fate in the retina.

                          (Chanut et al., 2000)

                          gro and Rpd3 appear to form a complex in vivo and interact directly via the glycine/proline rich (GP) domain.

                          (Chen et al., 1999)

                          gro mediated repression requires Rpd3 function.

                          (Chen et al., 1999)

                          tin protein interacts directly with the gro corepressor protein, for which the homeodomain is required.

                          (Choi et al., 1999)

                          gro protein mediates repressor activity of the hkb protein.

                          (Goldstein et al., 1999)

                          CtBP and gro repressors mediate separate pathways of transcriptional repression, and the two proteins can inhibit one another when both bind the same repressor.

                          (Zhang and Levine, 1999)

                          The products of the gro and pan genes interact to repress wg signalling activity.

                          (Cavallo et al., 1998)

                          run protein interacts with gro protein to repress transcription.

                          (Aronson et al., 1997)

                          Shows no genetic interaction with sdk.

                          (Nguyen et al., 1997)

                          Transcription complexes containing gro may associate with the amino terminus of His3 and these interactions may be propogated along the chromosomes due to the ability of gro to participate in higher order structures.

                          (Palaparti et al., 1997)

                          A screen to isolate mutations that cause dominant enhancement of wing vein phenotypes associated with Egfr recovered mutations in gro.

                          (Price et al., 1997)

                          The WRPW motif (the repressor domain) maps to the carboxy terminal of h-related proteins and is both necessary and sufficient to form protein-protein interactions with gro. gro is able to repress transcription when directly bound to DNA in the absence of binding to h-related proteins.

                          (Fisher et al., 1996)

                          Clones mutant for E(spl)-C bHLH-encoding genes or for gro display bristle hyperplasia. The E(spl)-C genes participate in the N signalling pathway. E(spl)-C mutants are epistatic over a gain of function mutant of N and ac-sc mutants are epistatic over E(spl)-C mutants.

                          (Heitzler et al., 1996)

                          Expression in Schneider cells demonstrates HLHm5 and E(spl) mediate transcriptional repression of an ac Ecol\CAT reporter gene, gro potentiates this effect.

                          (Heitzler et al., 1996)

                          The intracellular distribution and phosphorylation state of gro protein has been analysed.

                          (Husain et al., 1996)

                          E(spl) and gro form a highly interconnected interacting network involved in transcriptional regulation.

                          (Alifragis and Delidakis, 1995)

                          The bristle loss phenotype of H mutants can be suppressed by deleting components of the E(spl)-complex. The degree of suppression depends on both the number and identity of E(spl)-complex transcription units removed.

                          (Bang et al., 1995)

                          gro is crucial for transcriptional repression in several independent developmental pathways and is involved in establishing the terminal embryonic pattern.

                          (Ish-Horowicz et al., 1995)

                          gro and hh regulate en expression in the anterior compartment of the wing.

                          (de Celis and Ruiz-Gomez, 1995)

                          gro is required maternally for neurogenesis, segmentation and sex determination and acts as a transcriptional corepressor recruited to specific target promoters by h-related bHLH proteins.

                          (Paroush et al., 1994)

                          Arrangement and sequence of E(spl)-complex genes in D.melanogaster and D.hydei revealed that the E(spl)-gene, and the structure of complex are highly conserved, suggesting that each individual gene, as well as the organization of the complex, is of functional importance.

                          (Maier et al., 1993)

                          Individuals homozygous for Bpt1 and gro exhibit no Bpt1 phenotype.

                          (Duus et al., 1992)

                          gro is a neurogenic gene showing prominant maternal expression and is the only gene of the E(spl)-complex essential for viability. E(spl)-complex genes act as a functional unit composed of redundant genes which can partially substitute for each other. Eight E(spl)-region genes are required for the development of neurectodermal cells: HLHmδ, HLHmβ, HLHmγ, HLHm3, HLHm5, HLHm7, E(spl) and gro.

                          (Schrons et al., 1992)

                          gro allelic combinations show weak to intermediate neurogenic phenotypes.

                          (Delidakis et al., 1991)
                          Origin and Etymology
                          Discoverer
                          Etymology
                          Identification
                          External Crossreferences and Linkouts ( 95 )
                          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
                          Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
                          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
                          Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
                          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
                          Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                          GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
                          iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
                          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)
                          modMine - A data warehouse for the modENCODE project
                          SignaLink - A signaling pathway resource with multi-layered regulatory networks.
                          Linkouts
                          BioGRID - A database of protein and genetic interactions.
                          Cell Signaling Technology - Commercial vendor for primary antibodies and antibody conjugates.
                          Cell Signaling Technology - Commercial vendor for primary antibodies and antibody conjugates.
                          DroID - A comprehensive database of gene and protein interactions.
                          DRSC - Results frm RNAi screens
                          Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
                          FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                          FlyMine - An integrated database for Drosophila genomics
                          Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                          KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
                          MIST (genetic) - An integrated Molecular Interaction Database
                          MIST (protein-protein) - An integrated Molecular Interaction Database
                          Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                          Synonyms and Secondary IDs (30)
                          Reported As
                          Symbol Synonym
                          BEST:GM01575
                          BEST:LD15161
                          BcDNA.LD33829
                          (Rubin, 1999.4.23)
                          BcDNA:LD33829
                          CG8384
                          (Moskalev et al., 2018, Ni et al., 2011.7.19, Keleman et al., 2009.8.5, Bhutkar et al., 2008, Talamillo et al., 2008, Schlatter and Maier, 2005)
                          DGro
                          (Bajoghli, 2007)
                          E(spl)-WD
                          (Schmidt and Sladek, 1993)
                          E(spl)2
                          (Price et al., 1997)
                          E(spl)gro
                          (Bhattacharya and Baker, 2009, Li and Baker, 2001)
                          E(spl)m9/m10
                          (Delidakis et al., 1991)
                          En-spl
                          GRO
                          (Filion et al., 2010, van Steensel et al., 2010, Hittinger and Carroll, 2008, Choi et al., 2005, Taylor-Harding et al., 2004, Silver et al., 2003)
                          Grg/TLE
                          (Malin et al., 2005)
                          Gro
                          (Johnson, 2021, Gou et al., 2020, Nagel et al., 2020, Rust et al., 2020, Small and Arnosti, 2020, Smits and Shvartsman, 2020, Flint Brodsly et al., 2019, Miller et al., 2019, Steinmetz et al., 2019, Abed et al., 2018, Steinmetz et al., 2018, Chambers et al., 2017, Koltun et al., 2017, Zehender et al., 2017, Zhu et al., 2017, Jussen et al., 2016, Kwon et al., 2016, Auer et al., 2015, Lesnik et al., 2015, Jenny and Basler, 2014, Mannervik, 2014, Nagel and Preiss, 2014, Slattery et al., 2014, Zacharioudaki and Bray, 2014, Mbodj et al., 2013, Anderson et al., 2012, Holmqvist et al., 2012, Zarifi et al., 2012, Abed et al., 2011, Barry et al., 2011, Eastwood et al., 2011, Kim et al., 2011, Nagel and Preiss, 2011, Nègre et al., 2011, Parrott et al., 2011, Winkler et al., 2010, Chang et al., 2008, Kahali et al., 2008, Maier et al., 2008, Martinez and Arnosti, 2008, Winkler et al., 2008, Yao et al., 2008, Bejarano et al., 2007, Cinnamon et al., 2007, Tountas and Fortini, 2007, Von Ohlen et al., 2007, Moorman et al., 2006, Moorman et al., 2006, Stadeli, 2006, Wang and Jones, 2006, Kenyon et al., 2005, Kenyon et al., 2005, Ma, 2005, Nagel et al., 2005, Song et al., 2004, Wheeler et al., 2002)
                          Groucho
                          (Shukla et al., 2017, Holmqvist et al., 2012, Nedelsky et al., 2010)
                          anon-WO0118547.385
                          dAES1
                          (Chen and Courey, 2000)
                          dAES2
                          (Chen and Courey, 2000)
                          dGro
                          (Chen and Courey, 2000)
                          gro
                          (Martin et al., 2021, Pang et al., 2021, Trevino et al., 2021, Mira and Morante, 2020, Nair et al., 2020, Panta et al., 2020, Couturier et al., 2019, Guo et al., 2019, Shokri et al., 2019, Bischof et al., 2018, Davis and Rebay, 2018, Gene Disruption Project members, 2018-, Salazar and Yamamoto, 2018, Webber et al., 2018, Liu and Jin, 2017, Majot and Bidwai, 2017, Transgenic RNAi Project members, 2017-, van Tienen et al., 2017, Bivik et al., 2016, Johnston et al., 2016, Morimoto et al., 2016, Sarov et al., 2016, Baëza et al., 2015, Chen et al., 2015, Forés et al., 2015, Golubyatnikov et al., 2015, Kiparaki et al., 2015, Ashwal-Fluss et al., 2014, Boyle et al., 2014, Dewald et al., 2014, Rhee et al., 2014, Wurmbach and Preiss, 2014, Atkins et al., 2013, de Celis et al., 2013.9.11, Kwon et al., 2013, Sopko and Perrimon, 2013, Upadhyai and Campbell, 2013, Chen and Verheyen, 2012, Domanitskaya and Schüpbach, 2012, Formaz-Preston et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Marbach et al., 2012, Smith et al., 2012, Spokony and White, 2012.5.22, Technau et al., 2012, Turki-Judeh and Courey, 2012, Ajuria et al., 2011, Helman et al., 2011, Pérez et al., 2011, Toku et al., 2011, Walrad et al., 2011, Wang et al., 2011, Bardin et al., 2010, Walrad et al., 2010, Aerts et al., 2009, Babaoglan et al., 2009, Benítez et al., 2009, Bhattacharya and Baker, 2009, Cai and Laughon, 2009, Chanet et al., 2009, Christensen et al., 2009.2.28, Dworkin et al., 2009, Lee et al., 2009, Payankaulam and Arnosti, 2009, Salzer and Kumar, 2009, Stöbe et al., 2009, Zhai et al., 2009, Ziv et al., 2009, Bhutkar et al., 2008, Chang et al., 2008, Cinnamon et al., 2008, de Wit et al., 2008, Estella and Mann, 2008, Haussmann et al., 2008, Jennings et al., 2008, Kaspar et al., 2008, Lu et al., 2008, Mieszczanek et al., 2008, Protzer et al., 2008, Schmidt et al., 2008, Bejarano et al., 2007, Chao et al., 2007, Crews, 2007.1.6, Goodfellow et al., 2007, Liotta et al., 2007, Nagel et al., 2007, Orian et al., 2007, Payankaulam and Arnosti, 2007, Fang et al., 2006, Jennings et al., 2006, Jiang and Crews, 2006, Koelzer and Klein, 2006, Ratnaparkhi et al., 2006, Shirangi et al., 2006, Wehn and Campbell, 2006, Choi et al., 2005, Goldstein et al., 2005, Roederer et al., 2005, Schlatter and Maier, 2005, Gim et al., 2001, Lawrence et al., 2000)
                          groucho
                          (Chen et al., 2015, Rhee et al., 2014)
                          l(3)gro
                          (Nagel and Preiss, 1997, Ish-Horowicz et al., 1995)
                          m10
                          (Campos-Ortega and Knust, 1990, Knust et al., 1987)
                          m9
                          (Tata and Hartley, 1991, Campos-Ortega and Knust, 1990, Knust et al., 1987)
                          m9,m10
                          (Campos-Ortega and Knust, 1990)
                          m9-m10
                          (Campos-Ortega, 1993)
                          m9/10
                          (Muskavitch, 1994, Cornish et al., 1991, Marte et al., 1991, Preiss et al., 1991, Schrons et al., 1991)
                          m9/m10
                          (Schmidt and Sladek, 1993, Tietze et al., 1992, Cornish et al., 1990, Preiss et al., 1990, Klambt et al., 1989)
                          Name Synonyms
                          Dgroucho
                          (Brugmann et al., 2004)
                          Groucho
                          (Shi et al., 2020, Chambers et al., 2017, Zhu et al., 2017, Contreras-Cornejo et al., 2016, Jussen et al., 2016, Auer et al., 2015, Fiedler et al., 2015, Forés et al., 2015, Kaul et al., 2015, Dewald et al., 2014, Jenny and Basler, 2014, Nagel and Preiss, 2014, Bejsovec, 2013, Chen and Verheyen, 2012, Technau et al., 2012, Helman et al., 2011, Kim et al., 2011, Kurth et al., 2011, Parrott et al., 2011, Song et al., 2010, van Steensel et al., 2010, Walrad et al., 2010, Winkler et al., 2010, Arce et al., 2009, Cai and Laughon, 2009, Chen and Rasmuson-Lestander, 2009, Nie et al., 2009, Payankaulam and Arnosti, 2009, Syu et al., 2009, Von Ohlen and Moses, 2009, Cinnamon et al., 2008, Jennings et al., 2008, Kahali et al., 2008, Maier et al., 2008, Martinez and Arnosti, 2008, Talamillo et al., 2008, Winkler et al., 2008, Yao et al., 2008, Zhang et al., 2008, Bajoghli, 2007, Bejarano et al., 2007, Cinnamon et al., 2007, Gerlitz et al., 2007, Liotta et al., 2007, Metzger et al., 2007, Ohlen et al., 2007, Orian et al., 2007, Reig et al., 2007, Tountas and Fortini, 2007, He and Axelrod, 2006, Moorman et al., 2006, Parrish et al., 2006, Payankaulam et al., 2006, Shirangi et al., 2006, Stadeli, 2006, Wang and Jones, 2006, Goldstein et al., 2005, Malin et al., 2005, Nagel et al., 2005, Song et al., 2004, Jennings, 2002, Jia et al., 2002, Klochendler-Yeivin et al., 2002, Wheeler et al., 2002, Gim et al., 2001, Kenyon et al., 2001, Weinmaster, 1997)
                          Tle/Groucho
                          (Shi et al., 2020)
                          groucho
                          (Salazar and Yamamoto, 2018, Young and Marcotte, 2017, Golubyatnikov et al., 2015, Slattery et al., 2014, Formaz-Preston et al., 2012, Kuo et al., 2011, Bardin et al., 2010, Duncan and Dearden, 2010, Mummery-Widmer et al., 2009, Shevelyov et al., 2009, Zhai et al., 2009, González et al., 2008, Bejarano et al., 2007, Payankaulam and Arnosti, 2007, Uhler et al., 2007, Von Ohlen et al., 2007, Payankaulam et al., 2006, Wehn and Campbell, 2006, Choi et al., 2005, Roederer et al., 2005, Schlatter and Maier, 2005, Takanaka and Courey, 2005, Yu et al., 2005, Cowden and Levine, 2003, Silver et al., 2003, Lawrence et al., 2000, Stifani et al., 1992)
                          Secondary FlyBase IDs
                          • FBgn0019999
                          • FBgn0020008
                          • FBgn0023439
                          • FBgn0043006
                          • FBgn0043009
                          • FBgn0062049
                          Datasets (1)
                          Study focus (1)
                          Experimental Role
                          Project
                          Project Type
                          Title
                          • transgene_used
                          FBGSE22069
                          Protein profiling reveals five principal chromatin types in Drosophila cells.
                          Study result (0)
                          Result
                          Result Type
                          Title
                          References (533)