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General Information
Symbol
Dmel\Trl
Species
D. melanogaster
Name
Trithorax-like
Annotation Symbol
CG33261
Feature Type
FlyBase ID
FBgn0013263
Gene Model Status
Stock Availability
Gene Snapshot
Trithorax-like (Trl) encodes a GAGA transcription factor involved in chromatin modification. It contributes to cell division, dosage compensation and gametogenesis. [Date last reviewed: 2019-03-14]
Also Known As

GAF, GAGA, GAGA factor, Nc70F, GAGA-factor

Key Links
Genomic Location
Cytogenetic map
Sequence location
3L:14,747,929..14,761,049 [-]
Recombination map

3-42

RefSeq locus
NT_037436 REGION:14747929..14761049
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Summaries
Gene Group (FlyBase)
POLYCOMB GROUP RECRUITERS/DNA-BINDING PROTEINS -
The Polycomb group (PcG) proteins are epigenetic regulators, best characterized for the repression of Hox gene expression. In Drosophila, PcG proteins repress their target genes by binding to specific DNA elements called Polycomb Response Elements (PREs). PcG DNA binding proteins contribute to the recruitment of Polycomb complexes to PREs. (Adapted from FBrf0228921).
C2H2 ZINC FINGER TRANSCRIPTION FACTORS -
Zinc finger C2H2 transcription factors are sequence-specific DNA binding proteins that regulate transcription. They possess DNA-binding domains that are formed from repeated Cys2His2 zinc finger motifs. (Adapted from PMID:1835093, FBrf0220103 and FBrf0155739).
Protein Function (UniProtKB)
Transcriptional activator that functions by regulating chromatin structure. Overcomes the repressive effects of chromatin by promoting the open chromatin conformation in promoter gene regions, thereby allowing access to other transcription factors. Binds to DNA Polycomb response elements (PREs) at the bithorax complex and to the proximal region of the engrailed promoter, and positively regulates transcription of many genes including homeotic ones. Binds to the DNA sequence (GA)n, with optimal binding to the pentamer 5'-GAGAG-3'. Binds DNA as an oligomer. May also act as a transcriptional repressor, maintaining the repressed state of genes including lolal, and down-regulating its own transcription. Required for dosage compensation in males and may be involved in oogenesis. Also has a role in nuclear division.
(UniProt, Q08605)
Summary (Interactive Fly)

transcription factor - GAGA - Zinc finger - BTD domain - a chromatin modifying protein - cell division, dosage compensation and gametogenesis

Gene Model and Products
Number of Transcripts
10
Number of Unique Polypeptides
5

Please see the GBrowse view of Dmel\Trl or the JBrowse view of Dmel\Trl 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
Comments on Gene Model

gene_with_dicistronic_mRNA ; SO:0000722

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.

Gene model reviewed during 5.46

Gene model reviewed during 5.55

A non-AUG start codon may be used for translation of one or more transcripts of this gene; based on the presence of conserved protein signatures within the 5' UTR without an in-frame AUG (FBrf0243886).

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0075705
3197
582
FBtr0075707
4441
519
FBtr0075708
1977
519
FBtr0075706
4015
519
FBtr0075704
2156
519
FBtr0075703
2613
582
FBtr0100444
1911
519
FBtr0100445
3806
567
FBtr0333045
3814
608
FBtr0333046
3145
623
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0089417
62.6
582
8.68
FBpp0089419
54.8
519
8.13
FBpp0089411
54.8
519
8.13
FBpp0089418
54.8
519
8.13
FBpp0089416
54.8
519
8.13
FBpp0089415
62.6
582
8.68
FBpp0099866
54.8
519
8.13
FBpp0099867
60.4
567
9.55
FBpp0305259
64.2
608
8.67
FBpp0305260
66.4
623
7.72
Polypeptides with Identical Sequences

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

582 aa isoforms: Trl-PA, Trl-PF
519 aa isoforms: Trl-PB, Trl-PC, Trl-PD, Trl-PE, Trl-PH
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

Interacts with Bin1, lolal, corto, ttk and ph-p (PubMed:11256608, PubMed:12384587, PubMed:12834867, PubMed:12771214). Interacts with FACT subunits Ssrp and dre4/SPT16 (PubMed:12815073). Interacts with E(bx) (PubMed:11583616). Upon ecdysone stimulation, interacts with Nup98 (PubMed:28366641).

(UniProt, Q08605)
Post Translational Modification

The N-terminus is blocked.

(UniProt, Q08605)
Domain

The N-terminal BTB domain mediates protein oligomerization. The C-terminal glutamine-rich region is required for transcriptional activation activity.

(UniProt, Q08605)
Crossreferences
PDB - An information portal to biological macromolecular structures
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
Mapped Features

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

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (32 terms)
Molecular Function (11 terms)
Terms Based on Experimental Evidence (11 terms)
CV Term
Evidence
References
inferred from direct assay
inferred from mutant phenotype
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
inferred from mutant phenotype
(assigned by UniProt )
inferred from direct assay
inferred from physical interaction with UniProtKB:Q7KRI2
(assigned by UniProt )
inferred from physical interaction with FLYBASE:Ssrp; FB:FBgn0010278
inferred from physical interaction with UniProtKB:P17789
(assigned by UniProt )
inferred from physical interaction with UniProtKB:P41046
(assigned by UniProt )
inferred from physical interaction with UniProtKB:Q9VCH5
(assigned by UniProt )
inferred from physical interaction with UniProtKB:Q08605
(assigned by UniProt )
Terms Based on Predictions or Assertions (0 terms)
Biological Process (16 terms)
Terms Based on Experimental Evidence (14 terms)
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
(assigned by UniProt )
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from direct assay
(assigned by UniProt )
inferred from mutant phenotype
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
non-traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN001135010
(assigned by GO_Central )
Cellular Component (5 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN001135010
(assigned by GO_Central )
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: maternally deposited

Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

Trl protein is localized to the nuclei of all cell types in the pupal retina

Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
inferred from direct assay
(assigned by UniProt )
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Trl in GBrowse 2
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 4-6
  • Stages(s) 11-12
  • Stages(s) 13-16
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 76 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 24 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Trl
Transgenic constructs containing regulatory region of Trl
Deletions and Duplications ( 10 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
microchaeta & abdominal segment 6 | ectopic
mitosis & nuclear chromosome
mitosis & nuclear chromosome, with Scer\GAL4- :
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (18)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
3 of 15
Yes
Yes
1 of 15
No
Yes
1 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
Yes
1 of 15
No
Yes
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (13)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
No
1 of 15
Yes
Yes
1 of 15
Yes
No
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
No
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
No
1 of 15
Yes
Yes
Rattus norvegicus (Norway rat) (11)
1 of 13
Yes
Yes
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
Yes
1 of 13
Yes
No
1 of 13
Yes
Yes
Xenopus tropicalis (Western clawed frog) (9)
1 of 12
Yes
Yes
1 of 12
Yes
Yes
1 of 12
Yes
Yes
1 of 12
Yes
Yes
1 of 12
Yes
Yes
1 of 12
Yes
Yes
1 of 12
Yes
Yes
1 of 12
Yes
Yes
1 of 12
Yes
Yes
Danio rerio (Zebrafish) (6)
1 of 15
Yes
No
1 of 15
Yes
Yes
1 of 15
Yes
No
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
No
Caenorhabditis elegans (Nematode, roundworm) (1)
1 of 15
Yes
Yes
Arabidopsis thaliana (thale-cress) (2)
1 of 9
Yes
No
1 of 9
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG0919089I )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091507EK )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W07Q0 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Rhodnius prolixus
Kissing bug
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( None identified )
No non-Insect Arthropod orthologies identified
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
No non-Arthropod Metazoa orthologies identified
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (24)
3 of 10
3 of 10
3 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
1 of 10
1 of 10
1 of 10
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 ( 0 )
    Allele
    Disease
    Interaction
    References
    Disease Associations of Human Orthologs (via DIOPT v7.1 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?
    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

    Please see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    esyN Network Diagram
    esyN Network Key:
    Suppression
    Enhancement

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    External Data
    Subunit Structure (UniProtKB)
    Interacts with Bin1, lolal, corto, ttk and ph-p (PubMed:11256608, PubMed:12384587, PubMed:12834867, PubMed:12771214). Interacts with FACT subunits Ssrp and dre4/SPT16 (PubMed:12815073). Interacts with E(bx) (PubMed:11583616). Upon ecdysone stimulation, interacts with Nup98 (PubMed:28366641).
    (UniProt, Q08605 )
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    External Data
    Linkouts
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    3L
    Recombination map

    3-42

    Cytogenetic map
    Sequence location
    3L:14,747,929..14,761,049 [-]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    70F4-70F4
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    70F1-70F4
    (determined by in situ hybridisation)
    70F-70F
    (determined by in situ hybridisation)
    70F1-70F2
    (determined by in situ hybridisation)
    70E-70F
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (38)
    Genomic Clones (19)
    cDNA Clones (147)
     

    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 sequences
    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)
    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
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of

    Source for identity of: Trl CG9343

    Source for database merge of

    Source for merge of: Trl anon- EST:fe2E12

    Source for merge of: Trl Nc70F

    Additional comments

    This gene was originally designated as 'trithorax-like' ('Trl') residing in the left arm of the third chromosome, based on the observation that some mutant alleles could cause homeotic transformations like mutations in trx (FBrf0076985). However, this assignment is not supported by subsequent reports (FBrf0107412, FBrf0155709). Indeed, the homeotic effect is lost when the right arm of the third chromosome of the original alleles is replaced by a genetically marked WT chromosome (unpublished data). To avoid potential confusion, we refer to the gene as 'GAGA factor' ('Gaf').

    One or more of the processed transcripts for this gene contain(s) two non-overlapping open reading frames (ORFs). The non-overlapping ORFs are represented by Trl and CG42507.

    Annotation CG33260 has been re-extended to its original form (corresponding to the annotation originally named CG13470) in release 5.1 of the genome annotation. The 3' end of release 5.1 annotation CG33260 (including some of the CDS) overlaps the 5' UTR of the neighbouring release 5.1 Trl annotation which is on the same strand.

    Source for merge of Trl Nc70F was sequence comparison ( date:000202 ).

    Other Comments

    DNA-protein interactions: genome-wide binding profile assayed for Trl protein in 0-12 hr embryos; see mE1_TFBS_Trl collection report.

    DNA-protein interactions: genome-wide binding profile assayed for Trl 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).

    DNA-protein interactions: genome-wide binding profile assayed for Trl protein in 0-12 hr embryos. Individual protein-binding experiments listed under "Samples" at GEO: 16245 (http://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE16245).

    RNAi generated by PCR using primers directed to this gene causes a cell growth and viability phenotype when assayed in Kc167 and S2R+ cells.

    RNAi screen using dsRNA made from templates generated with primers directed against this gene causes a cell growth and viability phenotype when assayed in Kc167 and S2R+ cells.

    Trl is required for dosage compensation in males.

    Genetic interaction data suggest that Trl may have a role in control of homeotic gene activity.

    Trl protein forms oligomers both in vitro and in vivo. The formation of these oligomers, which exist in solution in the absence of DNA binding, requires the N-terminal POZ domain.

    The Trl POZ domain mediates strong co-operative binding to multiple Trl protein binding sites but inhibits binding to single sites. Trl protein oligomerisation thus increases binding specificity by selecting only promoters with multiple Trl protein binding sites.

    Trl- and Iswi-mediated disruption of the chromatin structure within the promoter region of ftz activates transcription on the chromatin template.

    Two major primary isoforms of Trl are identified, GAGA-519 and GAGA-581. The isoforms share an N-terminal protein-protein interaction domain and a DNA binding domain, but differ in their C-termini. Both isoforms exhibit a full spectrum of biological activities.

    Silencing activity of the iab-7PRE in the bithorax complex is dependent upon Trl.

    Hsf interacts directly with Trl, Trl stabilises Hsf binding to heat shock elements (HSEs).

    In tissue culture cells Trl protein is also bound at those PREs which contain Trl consensus-binding sites.

    The Trl gene product is required for a variety of developmental loci that contain GAGA binding sites in their upstream regulatory regions. The effects of mutation in Trl are consistent with a role in chromatin remodeling. Other defects suggest an additional, more global role in chromosome structure and function, related to the association of the GAGA protein with heterochromatic satellite sequences observed throughout the cell cycle.

    The single zinc finger domain from the Trl protein, along with a stretch of amino acids at the N terminus of the finger, is sufficient for specific DNA binding.

    The grh and Trl factors bind to the 11bp tor response element in the tll promoter, raising the possibility that they are involved in the regulation of tll expression. Trl may be required for early events in oogenesis.

    UV cross linking technique has been used to study the in vivo distribution of Trl protein on Hsp70 and Hsp26. Prior to heat shock Trl protein is associated with the promoter regions of the uninduced Hsp70 and Hsp26 genes. Upon heat shock induction Trl protein is recruited to their transcription units with its distribution coincident with that of RNA polymerase II.

    Analysis of transcription from Hsp26 promoter deletion constructs indicates that Trl mediates anti-repression of the Hsp26 promoter in extracts from unstressed embryos, while Hsf activates the Hsp26 promoter in extracts from heat shocked embryos.

    Nucleosome remodelling factor is composed of at least four polypeptides that act in concert with the Trl transcription factor at the Hsp70 promoter.

    Hsf and Trl can cause nucleosome rearrangements which may lead to a refinement of nucleosome positions, nucleosome remodeling is ATP-dependent.

    Removal of the POZ (poxvirus and zinc finger) domain increases DNA binding affinity of the chromosome.

    Trl belongs to the trithorax group of genes, is required for normal expression of homeotic genes, for example Ubx, and is a modifier of position effect variegation.

    Trl encodes the GAGA factor which is involved in the formation of an accessible chromatin structure at promoter sequences.

    Trl binds to tubulin element 2 (TE2) and the intron of αTub84B, but not to tubulin element 1 (TE1).

    Trl is associated with regions of heterochromatin throughout the cell cycle, probably via a direct interaction with a GA/CT rich subset of the highly repetitive DNA sequences found in heterochromatin. Trl can also associate with specific DNA sequences even when the DNA is highly compacted in the mitotic chromosomes.

    Introduction of Trl protein during or after nucleosome assembly in vitro results in disruption of nucleosome structure at the Hsp70Aa promoter. Hsp70Aa promoter deletion analysis reveals that disruption of a preassembled nucleosome requires three or four GA/CT elements, nucleosome displacement requires at least two GA/CT elements.

    Trl promotes chromatin rearrangement at heat shock loci.

    The Trl gene product is a multipurpose transcriptional activator which binds to (GA/CT)n sites in a host of promoters.

    Trl protein exists in multiple forms. Trl cDNA 5A activates transcription in transfected tissue culture cells in a binding site-dependent manner.

    Trl plays in role in the regulation of Ubx transcription.

    Trl counteracts His1-mediated repression of transcription.

    The Trl protein probably acts as a transcriptional antirepressor of the Kr gene.

    Trl binds to eve regulatory sequences in vitro.

    The Trl product activates Ubx transcription selectively in a reconstituted in vitro reaction.

    Origin and Etymology
    Discoverer
    Etymology
    Identification
    External Crossreferences and Linkouts ( 112 )
    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 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/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
    BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
    Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
    Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
    Flygut - An atlas of the Drosophila adult midgut
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    KEGG Genes - Molecular building blocks of life in the genomic space.
    modMine - A data warehouse for the modENCODE project
    PDB - An information portal to biological macromolecular structures
    SignaLink - A signaling pathway resource with multi-layered regulatory networks.
    Linkouts
    ApoDroso - Functional genomic database for photoreceptor development, survival and function
    BioGRID - A database of protein and genetic interactions.
    DPiM - Drosophila Protein interaction map
    DroID - A comprehensive database of gene and protein interactions.
    DRSC - Results frm RNAi screens
    FLIGHT - Cell culture data for RNAi and other high-throughput technologies
    FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
    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
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Synonyms and Secondary IDs (38)
    Reported As
    Symbol Synonym
    E(var)3-trl
    E(var)62
    GAF
    (Maksimenko et al., 2020, Erceg et al., 2019, Gutierrez-Perez et al., 2019, Kuhn et al., 2019, Kyrchanova et al., 2019, Vorobyeva and Mazina, 2019, Haines and Eisen, 2018, Kaye et al., 2018, Mourad and Cuvier, 2018, Rennie et al., 2018, Wang et al., 2018, Boija et al., 2017, Ciabrelli et al., 2017, Cleard et al., 2017, El-Sharnouby et al., 2017, Kaye et al., 2017, Lomaev et al., 2017, Mourad et al., 2017, Pascual-Garcia et al., 2017, Urban et al., 2017, Chaharbakhshi and Jemc, 2016, Duarte et al., 2016, Khoroshko et al., 2016, Astakhova et al., 2015, Erokhin et al., 2015, Fuda et al., 2015, Magbanua et al., 2015, Wolle et al., 2015, Ho et al., 2014, McElroy et al., 2014, Pascual-Garcia et al., 2014, Van Bortle et al., 2014, Enderle and McNeill, 2013, Kellner et al., 2013, Kwak et al., 2013, Li and Zhou, 2013, Li et al., 2013, Mason-Suares et al., 2013, Oh et al., 2013, Rubin and Green, 2013, Schuettengruber and Cavalli, 2013, Tariq et al., 2013, Agelopoulos et al., 2012, Bartoletti et al., 2012, Bayarmagnai et al., 2012, Feller et al., 2012, Guertin et al., 2012, Holmqvist et al., 2012, Sakoparnig et al., 2012, Agrawal et al., 2011, Bonchuk et al., 2011, Demakov et al., 2011, Islam et al., 2011, Kharchenko et al., 2011, Nègre et al., 2011, Rach et al., 2011, Sultana et al., 2011, Vatolina et al., 2011, Vatolina et al., 2011, Vogelmann et al., 2011, Deal et al., 2010, Filion et al., 2010, Guertin and Lis, 2010, Huen and Russell, 2010, modENCODE Consortium et al., 2010, Nègre et al., 2010, Schoborg and Labrador, 2010, The modENCODE Consortium, 2010, The modENCODE Consortium, 2010, van Bemmel et al., 2010, van Steensel et al., 2010, Ardehali et al., 2009, Bartkuhn et al., 2009, Chen et al., 2009, Schuettengruber et al., 2009, Shevelyov et al., 2009, Chopra et al., 2008, Hauenschild et al., 2008, Petesch and Lis, 2008, Golovnin et al., 2007, Schuettengruber et al., 2007, Schwartz and Pirrotta, 2007, Hill, 2006, Martinez et al., 2006, Muller and Kassis, 2006, Negre et al., 2006, Salvaing et al., 2006, Tolhuis et al., 2006, Zhao et al., 2006, Brown et al., 2005, Zhao et al., 2005, Lund and van Lohuizen, 2004, Melnikova et al., 2004, Smith et al., 2004, Bloyer et al., 2003, Ishii and Laemmli, 2003, Sun et al., 2003, van Steensel et al., 2003, Dejardin and Cavalli, 2002, Huang et al., 2002, Leibovitch et al., 2002, Maurange and Paro, 2002, Schwendemann and Lehmann, 2002, Bejarano et al., 2001, Busturia et al., 2001, Katokhin et al., 2001, Martienssen, 2001, Patterson, 2001, van Steensel et al., 2001, van Steensel et al., 2001, Henikoff and Vermaak, 2000, Agianian et al., 1999, Wilkins and Lis, 1999, Lis, 1998, Pile and Cartwright, 1998, Wilkins and Lis, 1998, Baricheva et al., 1997, Wilkins and Lis, 1997, Todorovic et al., 1996, Sandaltzopoulos et al., 1995, Wall et al., 1995)
    GAGA
    (Kwon et al., 2016, Negi et al., 2015, Dorogova et al., 2014, Slattery et al., 2014, Di Croce and Helin, 2013, Piñeyro et al., 2013, Kvon et al., 2012, Riddle et al., 2012, Aran-Guiu et al., 2010, Font-Burgada et al., 2008, Vaquero et al., 2008, Kahn et al., 2006, Bonet et al., 2005, Canudas et al., 2005, Moon et al., 2005, Wu et al., 2005, Lehmann et al., 2004, Pagans et al., 2004, Schweinsberg et al., 2004, Erhardt et al., 2003, Furriols and Casanova, 2003, Lu et al., 2003, Mahmoudi et al., 2003, Salvaing et al., 2003, Simms et al., 2003, Duncan, 2002, Kosoy et al., 2002, Mahmoudi et al., 2002, Poux et al., 2002, Volpi et al., 2002, West et al., 2002, Bloyer et al., 2001, Granok et al., 2001, Greenberg and Schedl, 2001, Leibovitch et al., 2001, Mishra et al., 2001, Poux et al., 2001, Van Steensel et al., 2001, Xiao et al., 2001, Deuring et al., 2000, Espinas et al., 2000, Farkas et al., 2000, Jones and Kadonaga, 2000, Read et al., 2000, Argiropoulos and Brock, 1999, Bloyer et al., 1999, Cryderman et al., 1999, Espinas et al., 1999, Horard et al., 1999, Katsani et al., 1999, van Lohuizen, 1999, Weiler and Wakimoto, 1999, Brock and Hodgson, 1998, Cavalli and Paro, 1998, Csink and Henikoff, 1998, Gdula et al., 1998, Huang et al., 1998, Jimenez-Garcia et al., 1998, Lehmann et al., 1998, Lintermann et al., 1998, Platero et al., 1998, Wu et al., 1998, Baricheva et al., 1997, Breiling et al., 1997, Greenberg and Schedl, 1997, Greenblatt, 1997, Platero et al., 1997, Schwartz and Kornberg, 1997, Strutt et al., 1997, Bhat et al., 1996, Ito et al., 1996, Laney and Biggin, 1996, Pedone et al., 1996, Pedone et al., 1996, Treisman, 1996, Yamamoto et al., 1996, Bienz and Muller, 1995, Hu et al., 1995, Liaw et al., 1995, O'Brien et al., 1995, Sandaltzopoulos and Bonte, 1995, Shopland et al., 1995, Wallrath and Elgin, 1995, Bardwell and Treisman, 1994, Lu et al., 1993, TenHarmsel et al., 1993, Laney and Biggin, 1992, Lee et al., 1992, Lu et al., 1992, Croston et al., 1991, Kerrigan et al., 1991, Elgin, 1990, Read et al., 1990, Thummel, 1989)
    TfGAGA/Adf-2
    Trl
    (Birnbaum et al., 2019, FlyBase Genome Annotators, 2019-, Shokri et al., 2019, Ueberschär et al., 2019, Bischof et al., 2018, Dasari et al., 2018, Kaye et al., 2018, Kyrchanova et al., 2018, Prange et al., 2018, Srivastava et al., 2018, Webber et al., 2018, Batut and Gingeras, 2017, Cleard et al., 2017, Houtz et al., 2017, Lee et al., 2017, Maini et al., 2017, Sharma et al., 2017, Transgenic RNAi Project members, 2017-, Yeung et al., 2017, Dorogova et al., 2016, Kwon et al., 2016, Quijano et al., 2016, Dietz et al., 2015, Gene Disruption Project members, 2015-, Ghasemi et al., 2015, model organism Encyclopedia of Regulatory Network (modERN) Project, 2015-, Naval-Sánchez et al., 2015, Oktaba et al., 2015, Schertel et al., 2015, Zabidi et al., 2015, Ashwal-Fluss et al., 2014, Boyle et al., 2014, Comoglio and Paro, 2014, Dorogova et al., 2014, Jiang and Singh, 2014, Rhee et al., 2014, Shlyueva et al., 2014, Aleksic et al., 2013, Darbo et al., 2013, Elgin and Reuter, 2013, Gisselbrecht et al., 2013, Karagodin et al., 2013, Lagha et al., 2013, Piñeyro et al., 2013, Vasanthi et al., 2013, Agelopoulos et al., 2012, Antao et al., 2012, Bartoletti et al., 2012, Bayarmagnai et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Kvon et al., 2012, Abruzzi et al., 2011, Carreira et al., 2011, Di Stefano et al., 2011, Fay et al., 2011, Nègre et al., 2011, Omelina et al., 2011, Pruteanu-Malinici et al., 2011, Rodriguez-Jato et al., 2011, Seong et al., 2011, Sultana et al., 2011, Toku et al., 2011, Baig et al., 2010, modENCODE Consortium et al., 2010, Saj et al., 2010, Smulders-Srinivasan et al., 2010, Vasanthi et al., 2010, Brody, 2009.03, Chen et al., 2009, Khan et al., 2009, Schaaf et al., 2009, Sing et al., 2009, Chopra et al., 2008, Dos-Santos et al., 2008, FlyBase Genome Annotators, 2008, Fujioka et al., 2008, González et al., 2008, Ogienko et al., 2008, Vázquez et al., 2008, Aerts et al., 2007, Bernués et al., 2007, Minidorff et al., 2007, Minidorff et al., 2007, Nakayama et al., 2007, Quinones-Coello, 2007, Ringrose and Paro, 2007, Secombe et al., 2007, Dworkin and Gibson, 2006, Dworkin and Gibson., 2006, Fedorova et al., 2006, Muller and Kassis, 2006, Bonet et al., 2005, Canudas et al., 2005, Tchoubrieva and Gibson, 2004, Balasov, 2002, Huang et al., 2002, Gim et al., 2001, Greenberg and Schedl, 2001, Hodgson et al., 2001, Katokhin et al., 2001, Trunova et al., 2001)
    anon-EST:fe2E12
    l(3)s2325
    Name Synonyms
    Adh transcription factor 2
    GAGA factor
    (Erceg et al., 2019, Webber et al., 2018, Cleard et al., 2017, Lomaev et al., 2017, Duarte et al., 2016, Tsai et al., 2016, Magbanua et al., 2015, Mason-Suares et al., 2013, Tariq et al., 2013, Agelopoulos et al., 2012, Bayarmagnai et al., 2012, Nakayama et al., 2012, Bonchuk et al., 2011, Islam et al., 2011, Vogelmann et al., 2011, Deal et al., 2010, Gurudatta et al., 2010, Huen and Russell, 2010, Kim et al., 2010, modENCODE Consortium et al., 2010, van Steensel et al., 2010, Vasanthi et al., 2010, Bartkuhn et al., 2009, Dhanasekaran et al., 2008, Kwong et al., 2008, Lee et al., 2008, Petesch and Lis, 2008, Bai et al., 2007, Dhanasekaran et al., 2007, Nakayama et al., 2007, Schuettengruber et al., 2007, Kahn et al., 2006, Moorman et al., 2006, Muller and Kassis, 2006, Negre et al., 2006, Tolhuis et al., 2006, Zhao et al., 2005, Fourel et al., 2004, Levine et al., 2004, Ringrose et al., 2004, Xin et al., 2004, Ishii and Laemmli, 2003, Kapetanaki et al., 2003, Mulholland et al., 2003, Orlando, 2003, Shimojima et al., 2003, Wu et al., 2003, Fourel et al., 2002, Leibovitch et al., 2002, Simon and Tamkun, 2002, Brock et al., 2001, Huckriede, 2001, Poux et al., 2001, Schweinsberg et al., 2001, Tchoubrieva and Gibson, 2001, Greenberg and Schedl, 2000, Horard et al., 2000, Leibovitch et al., 2000, Gao and Benyajati, 1998, Greenberg and Schedl, 1998, Klar, 1998, Okada and Hirose, 1998, Orlando et al., 1998, Schweinsberg et al., 1998, Anonymous, 1997, Benyajati et al., 1997, Leibovitch et al., 1997, Mason and Lis, 1997, Weber et al., 1997, Kingston et al., 1996, Marin et al., 1996, Kellum et al., 1995, Tsukiyama and Wu, 1995, Varga-Weisz et al., 1995, Wall et al., 1995, Becker, 1994, Granok et al., 1994, O'Donnell and Wensink, 1994, Kamakaka and Kadonaga, 1993, Krumm et al., 1993, Read and Manley, 1992, Biggin and Tjian, 1988)
    Neural conserved at 70F
    TRITHORAX-LIKE
    Trithorax-related
    anon-fast-evolving-2E12
    late boundary complex
    transcription-factor-Adh2
    Secondary FlyBase IDs
    • FBgn0036445
    • FBgn0053261
    • FBgn0005691
    • FBgn0010008
    • FBgn0010418
    • FBgn0011024
    • FBgn0025220
    Datasets (5)
    Study focus (5)
    Experimental Role
    Project
    Project Type
    Title
    • transgene_used
    The insulator protein SU(HW) modulates nuclear lamina interactions of the Drosophila genome.
    • transgene_used
    Protein profiling reveals five principal chromatin types in Drosophila cells.
    • bait_protein
    Genome-wide localization of chromosomal proteins in cell lines by ChIP-chip and ChIP-Seq.
    • bait_protein
    Genome-wide localization of chromosomal proteins in fly tissues by ChIP-chip and ChIP-Seq.
    • bait_protein
    Genome-wide localization of transcription factors by ChIP-chip and ChIP-Seq.
    References (572)