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General Information
Symbol
Dmel\l(2)gl
Species
D. melanogaster
Name
lethal (2) giant larvae
Annotation Symbol
CG2671
Feature Type
FlyBase ID
FBgn0002121
Gene Model Status
Stock Availability
Gene Snapshot
lethal (2) giant larvae (l(2)gl) encodes a tumor suppressor protein that regulates cell polarity and asymmetric cell division. It acts on the basolateral side of epithelial cells, antagonizing the activity of apical complex proteins encoded by baz, par-6 and aPKC. [Date last reviewed: 2019-09-26]
Also Known As
lgl, lethal giant larvae, p127, dlgl, p127l(2)gl
Key Links
Genomic Location
Cytogenetic map
Sequence location
2L:9,839..21,376 [-]
Recombination map
2-0
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
Belongs to the WD repeat L(2)GL family. (P08111)
Summaries
Pathway (FlyBase)
Negative Regulators of Notch Signaling Pathway -
The Notch receptor signaling pathway is activated by the binding of the transmembrane receptor Notch (N) to transmembrane ligands, Dl or Ser, presented on adjacent cells. This results in the proteolytic cleavage of N, releasing the intracellular domain (NICD). NICD translocates into the nucleus, interacting with Su(H) and mam to form a transcription complex, which up-regulates transcription of Notch-responsive genes. Negative regulators of the pathway down-regulate the signal from the sending cell or the response in the receiving cell. (Adapted from FBrf0225731 and FBrf0192604).
Protein Function (UniProtKB)
Essential for the development of polarized epithelia, for cell polarity associated with asymmetric cell division of neuroblasts during development, and for oocyte polarity formation. Promotes the formation of actin-rich projections at the oocyte cortex and the posterior enrichment of par-1 which is required for oocyte polarization. Regulates the localization of axis-specifying morphogens such as stau and grk. Isoform p78: Has an essential role in control of cell proliferation and differentiation during development and could act as a tumor suppressor. Isoform p127: Has an accessory function in control of cell proliferation and differentiation during development.
(UniProt, P08111)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
l(2)gl: lethal (2) giant larvae
Homozygotes undergo embryogenesis and the first three larval instars; larvae reach normal maximum size; then for some alleles most homozygotes fail to pupate, becoming bloated and 1.5-2 times normal size, whereas for others the majority form prepupae but fail to progress into morphogenesis. Ring gland small and appears immature in third-instar larvae (Scharrer and Hadorn, 1938, Proc. Nat. Acad. Sci. USA 24: 236-42); third-instar l(2)gl larvae implanted with a normal ring gland pupate but do not metamorphose; injection of ecdysone elicits the same result (Karlson and Hanser, 1952, Z. Naturforsch 76: 80-83); thus a deficiency of hormones from the ring gland is probably one, but not the only, result of l(2)gl. Homozygotes that die as prepupae have underdeveloped corpora allata and prothoracic glands, whereas larval lethals have underdeveloped prothoracic glands but normal corpora allata (Korochkina and Nazarova, 1977, Chromosoma 62: 175-90). Prothoracic glands contain approximately 1% the normal quantity of smooth endoplasmic reticulum (Aggarwal and King, 1969, J. Morph. 129: 171-99). Alleles range from 98% larval and 2% pupal death to 18% larval and 82% pupal death (Gateff, Golubovsky, and Sokolova, 1977, DIS 52: 128-29). In the most extreme phenotypes (+++), the larval brain and optic lobes become enlarged and disorganized and the imaginal discs large and clumped; when discs of such larvae are transplanted into wild-type-female abdomens, they form large contained tumors, whereas transplanted optic primordia from larval brains form invasive neuroblastomas, which grow rapidly, killing the host within 7-14 days; they can be serially cultured in adult abdomens (Gateff and Schneiderman, 1974). These observations have led to l(2)gl's being designated a Drosophila oncogene. Intermediate alleles (++) exhibit moderately enlarged brain and discs, which show enhanced growth when transplanted into wild-type females and death of host is delayed. In weak alleles (+) the brain and discs are small and rudimentary and grow slowly in transplants. One allele normal in disc morphology and behavior in transplants (-). The lethal phase is not well correlated with the phenotypic expression. Most abundant transcription noted in early (0-6 h) embryos and late third-instar larvae, with the smaller transcript more abundant in embryos and the larger in larvae (Mechler, McGinnis, and Gehring). Immunocytochemistry shows localization of l(2)gl product at cell surfaces, specifically at the interfaces between proliferating cells (Klambt and Schmidt, 1986, EMBO J. 5: 2955-61; Klambt, Muller, Lutzelschwab, Rossa, Totzke, and Schmidt, 1989, Dev. Biol. 133: 425-36; Lutzelschwab, Klambt, Rossa, and Schmidt, 1987, EMBO J. 6: 1791-97). During later embryogenesis relatively high amounts of l(2)gl protein is detected in pole cells and cells of the developing nervous system; specifically neurons in the peripheral nervous system undergoing axogenesis express the protein. Monoclonal antibodies specifically stain junctions between mammalian cells in culture as though they are recognizing either membrane or intercellular-matrix proteins (Klambt et al., 1989). Ten-to-eleven-day-old larvae homozygous for larval-lethal alleles exhibit remarkably few puff sites, only 63BC and occasionally 88D and 89B; however, heat shock induced puffs develop normally (Ashburner, 1970, Chromosoma 31: 356-76). Homozygotes able to form prepupae exhibit more nearly normal puffing patterns; puffing in response to administration of ecdysone also appears normal (Richards, 1976, Wilhelm Roux's Arch. Dev. Biol. 179: 339-48).
Summary (Interactive Fly)
novel conserved cytoskeletal element - functions with other tumor suppressors to regulate cell polarity and growth
Gene Model and Products
Number of Transcripts
11
Number of Unique Polypeptides
4

Please see the GBrowse view of Dmel\l(2)gl or the JBrowse view of Dmel\l(2)gl 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 model reviewed during 5.44
Annotated transcripts do not represent all supported alternative splices within 5' UTR.
Annotated transcripts do not represent all possible combinations of alternative exons and/or alternative promoters.
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.45
Multiphase exon postulated: exon reading frame differs in alternative transcripts.
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0078171
5391
1161
FBtr0078166
5155
1153
FBtr0078170
5248
1161
FBtr0078167
5230
1112
FBtr0078168
5225
1112
FBtr0078169
5082
1112
FBtr0306589
5315
1161
FBtr0306590
5179
1161
FBtr0306591
5390
1161
FBtr0306592
5528
1161
FBtr0330655
5281
1093
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
FBpp0077829
126.9
1161
5.78
FBpp0077824
125.9
1153
5.57
FBpp0077828
126.9
1161
5.78
FBpp0077825
121.2
1112
5.15
FBpp0077826
121.2
1112
5.15
FBpp0077827
121.2
1112
5.15
FBpp0297544
126.9
1161
5.78
FBpp0297545
126.9
1161
5.78
FBpp0297546
126.9
1161
5.78
FBpp0297547
126.9
1161
5.78
FBpp0303505
119.7
1093
6.01
Polypeptides with Identical Sequences

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

1112 aa isoforms: l(2)gl-PD, l(2)gl-PE, l(2)gl-PF
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)
May form multimeric complexes. Interacts with mahj. Interacts with aPKC; the interaction results in phosphorylation of l(2)gl.
(UniProt, P08111)
Post Translational Modification
Phosphorylated by aPKC which restricts l(2)gl activity to the oocyte posterior and is required for oocyte polarity formation.
(UniProt, P08111)
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\l(2)gl 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 (65 terms)
Molecular Function (7 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
inferred from genetic interaction with FLYBASE:zip; FB:FBgn0265434
inferred from genetic interaction with FLYBASE:zip; FB:FBgn0265434
inferred from physical interaction with MGI:MGI:893577
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000028271
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000028271
(assigned by GO_Central )
traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000028271
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000028271
(assigned by GO_Central )
Biological Process (45 terms)
Terms Based on Experimental Evidence (29 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:N; FB:FBgn0004647
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (18 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000812995
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000812995
(assigned by GO_Central )
non-traceable author statement
non-traceable author statement
non-traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000812995
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000812995
(assigned by GO_Central )
traceable author statement
non-traceable author statement
Cellular Component (13 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (9 terms)
CV Term
Evidence
References
traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000812995
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN000028271
(assigned by GO_Central )
non-traceable author statement
traceable author statement
traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000028271
(assigned by GO_Central )
non-traceable author statement
traceable author statement
non-traceable author statement
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
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
cell fractionation
Stage
Tissue/Position (including subcellular localization)
Reference
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
The l(2)gl protein is localized at cellular membranes or the intercellular matrix.
The l(2)gl protein is found in all primordia of larval tissues in the embryo. Neural primordia are only weakly labelled with the exception of the presumptive optic centers and axon bundles of the ventral cord which are distinctly labelled. In larvae, l(2)gl protein is seen at low levels until the end of the third instar where it is found mainly in brain and imaginal discs.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\l(2)gl 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
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 70 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 48 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of l(2)gl
Transgenic constructs containing regulatory region of l(2)gl
Deletions and Duplications ( 65 )
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
dorsal mesothoracic disc & epithelial cell
larval brain & neuroblast | supernumerary
larval brain | posterior & neuroblast | supernumerary
larval brain | posterior & neuron
microtubule & oocyte, with Scer\GAL4VP16.mat.αTub67C
microtubule & oocyte | germ-line clone
nurse cell & cytoskeleton | conditional ts
oocyte & cytoskeleton | conditional ts
ventral thoracic disc & epithelial cell
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (4)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
15 of 15
Yes
Yes
 
 
12 of 15
Yes
No
1 of 15
No
No
1 of 15
No
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (4)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
14 of 15
Yes
Yes
12 of 15
No
Yes
1 of 15
No
No
1 of 15
No
No
Rattus norvegicus (Norway rat) (4)
13 of 13
Yes
Yes
12 of 13
No
Yes
2 of 13
No
No
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (5)
8 of 12
Yes
Yes
7 of 12
No
Yes
1 of 12
No
Yes
1 of 12
No
No
1 of 12
No
Yes
Danio rerio (Zebrafish) (6)
12 of 15
Yes
Yes
12 of 15
Yes
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (3)
6 of 15
Yes
Yes
2 of 15
No
No
1 of 15
No
Yes
Arabidopsis thaliana (thale-cress) (2)
3 of 9
Yes
No
2 of 9
No
No
Saccharomyces cerevisiae (Brewer's yeast) (2)
4 of 15
Yes
No
4 of 15
Yes
No
Schizosaccharomyces pombe (Fission yeast) (1)
3 of 12
Yes
No
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091901TD )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091500OB )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
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) ( EOG090W00IY )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
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
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X00I4 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G01L0 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (1)
4 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 3 )
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 1 )
Allele
Disease
Interaction
References
Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
 
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.
Dmel gene
Ortholog showing functional complementation
Supporting References
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)
May form multimeric complexes. Interacts with mahj. Interacts with aPKC; the interaction results in phosphorylation of l(2)gl.
(UniProt, P08111 )
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
Gene Group - Pathway Membership (FlyBase)
Negative Regulators of Notch Signaling Pathway -
The Notch receptor signaling pathway is activated by the binding of the transmembrane receptor Notch (N) to transmembrane ligands, Dl or Ser, presented on adjacent cells. This results in the proteolytic cleavage of N, releasing the intracellular domain (NICD). NICD translocates into the nucleus, interacting with Su(H) and mam to form a transcription complex, which up-regulates transcription of Notch-responsive genes. Negative regulators of the pathway down-regulate the signal from the sending cell or the response in the receiving cell. (Adapted from FBrf0225731 and FBrf0192604).
External Data
Linkouts
KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
Genomic Location and Detailed Mapping Data
Chromosome (arm)
2L
Recombination map
2-0
Cytogenetic map
Sequence location
2L:9,839..21,376 [-]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
21A5-21A5
Limits computationally determined from genome sequence between P{lacW}spenk06805&P{lacW}l(2)k13604k13604 and P{lacW}U2af38k14504
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
21A-21A
(determined by in situ hybridisation)
21A3-21A4
(determined by in situ hybridisation)
Distal-most single copy gene of 2L.
Determined by comparing Celera genomic sequence with sequence from BDGP BAC and P1 clones.
Experimentally Determined Recombination Data
Location
Left of (cM)
Notes
Maps close to al.
Most distal gene on 2L.
Stocks and Reagents
Stocks (27)
Genomic Clones (5)
 

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

cDNA Clones (97)
 

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
 
polyclonal antibody, raised against the C-terminal peptide 1146-DNKIGTPKTAPEESQF-1161
Commercially Available Antibodies
 
Other Information
Relationship to Other Genes
Source for database identify of
Source for identity of: l(2)gl CG2671
Source for database merge of
Additional comments
A high incidence of mutant alleles found in natural populations in the Soviet Union (Golubovsky and Sokolova, 1973). Most tested alleles are molecular deficiencies with one breakpoint distal to the origin of the 40 kb walk and the other as indicated in allele list. Only l(2)gl52, which has a 10 kb insertion, and l(2)gl275, in which 7.9 kb have been deleted and replaced by an insert of 6.5 kb, are exceptions.
Other Comments
Second chromosome stocks often contain second-site deletions uncovering l(2)gl, the high frequency probably being due to the fact that l(2)gl is the second protein coding gene downstream of the subtelomeric region of chromosome 2L.
l(2)gl-induced metastases express both neuronal (elav) and glial cell (repo) type markers.
Loss of l(2)gl in larvae causes neoplastic brain tumors. Fragments of brat brain tumours transplanted into adult hosts over-proliferate and kill their hosts within 2 weeks.
Overexpression of l(2)gl in D.melanogaster males results in paternal-effect lethality that mimics the fertilisation defects associated with cytoplasmic incompatibility (CI) caused by Wolbachia infection.
The l(2)gl product regulates the membrane localization of the spdo protein.
l(2)gl promotes cortical localisation of mira.
Phosphorylation of l(2)gl by aPKC at the apical neuroblast cortex restricts l(2)gl activity and mira localisation to the opposite, basal side of the cell in embryonic neuroblasts.
l(2)gl acts together with numb in N inhibition and cell fate specification in the adult sensory organ precursor lineage.
dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.
dsRNA made from templates generated with primers directed against this gene is tested in an RNAi screen for effects on actin-based lamella formation.
l(2)gl is required downstream of dpp for the specification of dorsal epidermis.
The l(2)gl and dlg1 products act in a common process that differentially mediates cortical protein targeting in mitotic neuroblasts, creating intrinsic differences between daughter cells.
l(2)gl protein is uniformly cortical and interact with several types of myosin to localise neuroblast fate determinants.
The l(2)gl and dlg1 gene products regulate basal protein targeting, but not apical complex formation or spindle orientation, in both embryonic and larval neuroblasts. The l(2)gl and dlg1 proteins promote, and that of zip inhibits, actomyosin dependent basal protein targeting in neuroblasts.
l(2)gl plays a critical role at the onset of vitellogenesis and regulates growth of the oocyte, follicle cell migration and organisation and germline cell viability.
S.cerevisiae Scer\SRO7 Scer\SRO77 double mutants are partially functionally complemented by l(2)gl. S.cerevisiae Scer\SRO7 single mutants show slight complementation by l(2)gl.
Genetic analysis of l(2)gl reveal function is required during embryonic and post-embryonic development to maintain the normal developmental capacity.
Some of the proteins of apico-lateral junctions are required both for apico-basal cell polarity and for the signalling mechanisms controlling cell proliferation, whereas others are required more specifically in cell-cell signalling.
l(2)gl function is required for proper development during early embryogenesis.
p127 protein of l(2)gl is able to build quaternary structures forming a network with which other proteins associate. As revealed by the tumorous phenotype, organisation of the p127 network and its association with other proteins plays critical roles in the control of cell proliferation.
A serine kinase is tightly associated with l(2)gl. Activation of the serine kinase results in the disassociation of zip from the l(2)gl complex without affecting the homo-oligomerisation of l(2)gl.
The l(2)gl product is required in vivo in different types of epithelial cells to control their shape during development.
l(2)gl encodes a cytoskeletal protein that is required in aspects of Abl-dependent axonogenesis.
Mosaic analysis of l(2)gl mutants suggests a role for l(2)gl in cell-cell signaling and interactions.
The p127 protein of the l(2)gl gene is a component of a cytoskeletal network when complexed with a nonmuscle myosin II heavy chain protein, zip. Partial disruption of this complex causes l(2)gl gene inactivation and neoplastic transformation.
l(2)gl p127 protein is a component of a cytoskeletal network extending in the cytoplasm and/or underlaying the inner face of the plasma membrane in a variety of cells and tissues.
Mutants display a brain and imaginal disc neoplastic phenotype.
Mutations in l(2)gl cause malignant tumours in the brain and imaginal discs and generate developmental defects in a number of other tissues. Cellular and subcellular localisation of the protein to the cytoplasm and the inner face of the lateral cell membrane suggests that changes in cell shape and the loss of apical-basal polarity observed in tumorous tissues are a direct result of alterations in the cytoskeleton organisation caused by l(2)gl inactivation. Results also suggest the protein is involved in a cytoskeletal-based intercellular communication system directing cell differentiation.
The structure of the cytosolic form of l(2)gl protein confirms that the protein is a component of the cytoskeletal network including myosin and suggests that the neoplastic transformation resulting from l(2)gl gene inactivation may be caused by the partial disruption of this network.
When l(2)gl mutant brains are transplanted into wild-type adult hosts they can develop into enormous tumors. Using antibodies against the human 72kD type IV collagenase (differentially expressed in metastatic tumors), a cross reacting gelatinase of 49kD has been identified which is increased in l(2)gl mutant vs wild-type brains. Tumor cells that invade host tissues express Gelatinase, suggesting that the metastasis of Drosophila cells is similar to the metastasis of some human tumors at the biochemical as well as the cellular level.
awd function is required for brain tumour formation or proliferation in l(2)gl mutants. Mutant l(2)gl induced neuroblastomas are invasive and mutants have an increased proportion of awd expressing cells in the brain.
The highly divergent cis-regulatory elements of Dpse\l(2)gl can be fully recognized in D.melanogaster and lead to the synthesis of a transgenic protein that has enough specificity conserved for replacing the tumor-suppressor function normally fulfilled by the D.melanogaster l(2)gl protein.
Fas3, mys, disco, zip, l(2)gl, N and Egfr mutants show an additive phenotype in combination with Fas1TE89Da.
Neoplastic growth takes place in clones of cells that have lost l(2)gl in the preblastoderm syncytial embryos prior to any l(2)gl expression. Clones produced at the embryonic stages do not display the neoplastic phenotype and clones that arise in the larval stages show near normal or normal development. This analysis demonstrates the critical period for the establishment of tumorigenesis occurs during early embryogenesis at a time when l(2)gl expression is most intense in all cells. P-element transformation of l(2)gl deletion derivatives identify the essential domains.
The transcription patterns of Abl, R, Ras85D and Src64B were analyzed in neuroblasts derived from tumerous larval brain of l(2)gl larvae and S2 tissue culture cells.
Monoclonal antibodies have been raised against the l(2)gl protein and distribution patterns demonstrate that the l(2)gl protein is involved in proliferation arrest of cells.
l(2)gl has been cloned and the temporal pattern of RNA expression analysed.
The phenotype of a number of homozygous and transheterozygous l(2)gl mutants has been studied.
The viability of 7 l(2)gl alleles in heterozygotes has been studied at different temperatures. At 25oC, viability of the heterozygotes is reduced, but at low (12, 17oC) and high (29-30oC) temperatures the heterozygotes have a considerable advantage compared to wild-type.
Homozygotes undergo embryogenesis and the first three larval instars; larvae reach normal maximum size; then for some alleles most homozygotes fail to pupate, becoming bloated and 1.5-2 times normal size, whereas for others the majority form prepupae but fail to progress into morphogenesis. Ring gland small and appears immature in third instar larvae (Scharrer and Hadorn, 1938); third instar l(2)gl larvae implanted with a normal ring gland pupate but do not metamorphose; injection of ecdysone elicits the same result (Karlson and Hanser, 1952); thus a deficiency of hormones from the ring gland is probably one, but not the only, result of l(2)gl. Homozygotes that die as prepupae have underdeveloped corpora allata and prothoracic glands, whereas larval lethals have underdeveloped prothoracic glands but normal corpora allata (Korochkina and Nazarova, 1977). Prothoracic glands contain approximately 1% the normal quantity of smooth endoplasmic reticulum (Aggarwal and King, 1969). Alleles range from 98% larval and 2% pupal death to 18% larval and 82% pupal death (Gateff, Golubovsky and Sokolova, 1977). In the most extreme phenotypes, the larval brain and optic lobes become enlarged and disorganized and the imaginal discs large and clumped; when discs of such larvae are transplanted into wild-type-female abdomens, they form large contained tumors, whereas transplanted optic primordia from larval brains form invasive neuroblastomas, which grow rapidly, killing the host within 7-14 days; they can be serially cultured in adult abdomens (Gateff and Schneiderman, 1974). These observations have led to l(2)gl's being designated a Drosophila oncogene. Intermediate alleles exhibit moderately enlarged brain and discs, which show enhanced growth when transplanted into wild-type females and death of host is delayed. In weak alleles the brain and discs are small and rudimentary and grow slowly in transplants. One allele (l(2)gl558) normal in disc morphology and behavior in transplants. The lethal phase is not well correlated with the phenotypic expression. Most abundant transcription noted in early (0-6 h) embryos and late third instar larvae, with the smaller transcript more abundant in embryos and the larger in larvae (Mechler, McGinnis and Gehring, 1985). Immunocytochemistry shows localization of l(2)gl product at cell surfaces, specifically at the interfaces between proliferating cells (Klambt and Schmidt, 1986; Klambt, Muller, Lutzelschwab, Rossa, Totzke and Schmid; Lutzelschwab, Klambt, Rossa and Schmidt, 1987). During later embryogenesis relatively high amounts of l(2)gl protein is detected in pole cells and cells of the developing nervous system; specifically neurons in the peripheral nervous system undergoing axogenesis express the protein. Monoclonal antibodies specifically stain junctions between mammalian cells in culture as though they are recognizing either membrane or intercellular-matrix proteins (Klambt et al., 1989). Ten-to-eleven-day-old larvae homozygous for larval-lethal alleles exhibit remarkably few puff sites, only 63BC and occasionally 88D and 89B; however, heat shock induced puffs develop normally (Ashburner, 1970). Homozygotes able to form prepupae exhibit more nearly normal puffing patterns; puffing in response to administration of ecdysone also appears normal (Richards, 1976).
Origin and Etymology
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Etymology
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External Crossreferences and Linkouts ( 77 )
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.
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
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FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
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KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
MIST (genetic) - An integrated Molecular Interaction Database
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Synonyms and Secondary IDs (39)
Reported As
Symbol Synonym
Lgl
(Bonello and Peifer, 2019, Carrasco-Rando et al., 2019, Xu et al., 2019, Kon, 2018, Portela et al., 2018, Schmidt and Grosshans, 2018, Furuse and Izumi, 2017, Lang and Munro, 2017, Paglia et al., 2017, Richardson and Portela, 2017, Ajduk and Zernicka-Goetz, 2016, Calero-Cuenca et al., 2016, Clavería and Torres, 2016, Di Gregorio et al., 2016, Flores-Benitez and Knust, 2016, Izumi et al., 2016, Roman-Fernandez and Bryant, 2016, Schimizzi et al., 2016, Soriano et al., 2016, Zhang et al., 2016, Bajaj et al., 2015, Bell et al., 2015, Cao et al., 2015, Carvalho et al., 2015, Dong et al., 2015, Irvine and Harvey, 2015, Ohno et al., 2015, Portela et al., 2015, Schweisguth, 2015, Wang et al., 2015, Amoyel and Bach, 2014, Graybill and Prehoda, 2014, Jiang et al., 2014, Robbins et al., 2014, Rudrapatna et al., 2014, Wang et al., 2014, Bergstralh et al., 2013, Chen and Zhang, 2013, Enderle and McNeill, 2013, Goh et al., 2013, Khan et al., 2013, Levayer and Moreno, 2013, Noatynska et al., 2013, Yu and Guan, 2013, Fausti et al., 2012, Guilgur et al., 2012, Kelsom and Lu, 2012, Liu et al., 2012, Muñoz-Soriano et al., 2012, Neumüller et al., 2012, Tepass, 2012, Baker, 2011, Genevet and Tapon, 2011, Halder and Johnson, 2011, Hogan et al., 2011, Laprise and Tepass, 2011, Miles et al., 2011, Nance and Zallen, 2011, Zhao et al., 2011, Doerflinger et al., 2010, Duchi et al., 2010, Tamori et al., 2010, Yan, 2010, Coumailleau et al., 2009, Tian and Deng, 2009, Woolworth et al., 2009, Denef et al., 2008, Erben et al., 2008, Gervais et al., 2008, Kloepper et al., 2008, Kloepper et al., 2008, Leibfried et al., 2008, Grifoni et al., 2007, Grifoni et al., 2007, Leibfried et al., 2007, Tian and Deng, 2007, Wirtz-Peitz et al., 2007, Zarnescu et al., 2007, He and Axelrod, 2006, Qian and Prehoda, 2006, Solecki et al., 2006, Suzuki and Ohno, 2006, Swanson and Beitel, 2006, Szafranski and Goode, 2006, Wang et al., 2006, Betschinger et al., 2005, Budde, 2005, Budde, 2005, Dollar et al., 2005, Gangar et al., 2005, Georgiou et al., 2005, Langevin et al., 2005, Le Bivic, 2005, Mayer et al., 2005, Sanchez-Soriano et al., 2005, Somers and Chia, 2005, Wiggin et al., 2005, Macara, 2004, Wei et al., 2004, Wu and Beitel, 2004, Barros et al., 2003, Betschinger et al., 2003, Bilder, 2003, Goode and Szafranski, 2003, Henrique and Schweisguth, 2003, Hutterer et al., 2003, Kiehart, 2003, Knoblich et al., 2003, Müller and Bossinger, 2003, Rogers et al., 2003, Rolls et al., 2003, Tuxworth and Chia, 2003, Knust and Bossinger, 2002, Bilder, 2001, Leventis et al., 2001, Ohshiro et al., 2000, Peng et al., 2000)
MENE (2L)-B
l(2) giant larva
l(2)giant larvae
lgl
(Kurelac et al., 2019, Mirzoyan et al., 2019, Moreira et al., 2019, Sun et al., 2019, Daniel et al., 2018, Enomoto et al., 2018, Gerlach et al., 2018, Paul et al., 2018, Richardson and Portela, 2018, Banerjee and Roy, 2017, Ma et al., 2017, Ren et al., 2017, Shu et al., 2017, Tamori and Deng, 2017, Yasugi et al., 2017, Bergstralh et al., 2016, Calleja et al., 2016, Casas-Tintó et al., 2016, Johnson et al., 2016, Ma et al., 2016, Moreira and Morais-de-Sá, 2016, Bell et al., 2015, Besson et al., 2015, Grifoni et al., 2015, Sinkovics, 2015, Zhang et al., 2015, Berns et al., 2014, Johnston, 2014, Jones and Srivastava, 2014, Ma et al., 2014, Morais-de-Sá et al., 2014, Parsons et al., 2014, Wang et al., 2014, Bangi, 2013, Goh et al., 2013, Grifoni et al., 2013, Harvey et al., 2013, Hombría and Serras, 2013, Jagut et al., 2013, Levayer and Moreno, 2013, Ma et al., 2013, Ma et al., 2013, Staples and Broadie, 2013, Tamori and Deng, 2013, Carney et al., 2012, Fletcher et al., 2012, Haenfler et al., 2012, Izumi et al., 2012, Justiniano et al., 2012, Mirth and Shingleton, 2012, Song and Lu, 2012, Jukam and Desplan, 2011, Letizia et al., 2011, Li et al., 2011, Miles et al., 2011, Reddy and Irvine, 2011, Bahri et al., 2010, Bina et al., 2010, Chang et al., 2010, Colosimo et al., 2010, Fichelson et al., 2010, Froldi et al., 2010, Grzeschik et al., 2010, Grzeschik et al., 2010, Janic et al., 2010, Kaplan and Tolwinski, 2010, Krahn et al., 2010, Laprise et al., 2010, Menéndez et al., 2010, Menshchikova et al., 2010, Omelyanchuk and Pertseva, 2010, Portela et al., 2010, Robinson et al., 2010, Simone and DiNardo, 2010, Tong et al., 2010, Walther and Pichaud, 2010, Wu et al., 2010, Zeng et al., 2010, Atwood and Prehoda, 2009, Courbard et al., 2009, Gilbert et al., 2009, Gilbert et al., 2009, Huang et al., 2009, Kaplan et al., 2009, Li et al., 2009, Mao and Freeman, 2009, Ogawa et al., 2009, Parrish et al., 2009, Rhiner et al., 2009, Roegiers et al., 2009, Sousa-Nunes et al., 2009, Wang et al., 2009, Yamamoto et al., 2009.2.25, Bowman et al., 2008, Castellanos et al., 2008, Chia et al., 2008, Georgiou et al., 2008, Li et al., 2008, Pastor-Pareja et al., 2008, Tian and Deng, 2008, Vaccari et al., 2008, Weisman and Golubovsky, 2008, Wirtz-Peitz et al., 2008, Wirtz-Peitz et al., 2008, Yamamoto et al., 2008, Zhao et al., 2008, Beaucher et al., 2007, Beaucher et al., 2007, Blankenship et al., 2007, Grifoni et al., 2007, Grzeschik et al., 2007, Grzeschik et al., 2007, Guthridge et al., 2007, Menut et al., 2007, Slack et al., 2007, Srivastava et al., 2007, Szafranski and Goode, 2007, Tountas and Fortini, 2007, Tyler et al., 2007, Zarnescu et al., 2007, Betschinger et al., 2006, Denef et al., 2006, Humbert et al., 2006, Igaki et al., 2006, Igaki et al., 2006, Laprise et al., 2006, Lee et al., 2006, Lee et al., 2006, Pinter and Zarnescu, 2006, Wirtz-Peitz and Knoblich, 2006, Zhao et al., 2006, Beaucher et al., 2005, Broadie and Pan, 2005, Djiane et al., 2005, Goode et al., 2005, Pal et al., 2005, Richardson et al., 2005, Roegiers et al., 2005, Amin et al., 2004, Bardin et al., 2004, Beaucher et al., 2004, Betschinger and Knoblich, 2004, Bilder, 2004, Coffman, 2004, Grifoni et al., 2004, Grifoni et al., 2004, Hutterer et al., 2004, Lee et al., 2004, Llimargas et al., 2004, Roegiers and Jan, 2004, Szafranski and Goode, 2004, Abdelilah-Seyfried et al., 2003, Albertson and Doe, 2003, Bach et al., 2003, Beaucher et al., 2003, Bilder et al., 2003, Cereijido et al., 2003, Dow et al., 2003, Gibson and Perrimon, 2003, Grifoni et al., 2003, Horton and Ehlers, 2003, Humbert et al., 2003, Johnson and Wodarz, 2003, Justice and Jan, 2003, Justice et al., 2003, Muller, 2003, Nelson, 2003, Pagliarini and Xu, 2003, Petritsch et al., 2003, Roegiers et al., 2003, Rolls et al., 2003, Tanentzapf and Tepass, 2003, Wodarz and Huttner, 2003, Caruana, 2002, Chia and Yang, 2002, Johnston and Gallant, 2002, Medina et al., 2002, Wedlich, 2002, Bilder, 2001, Deng et al., 2001, Doe, 2001, Doe and Bowerman, 2001, Goodliffe et al., 2001, Greaves, 2001, Luxenberg et al., 2001, Ohshiro et al., 2001, Schaefer and Knoblich, 2001, Tanentzapf and Tepass, 2001, Tepass et al., 2001, Zelhof et al., 2001, Bilder et al., 2000, Greaves, 2000, Muller, 2000, Ohshiro et al., 2000, Peifer, 2000, Peng et al., 2000, Wodarz, 2000, Zahraoui et al., 2000, Bilder and Perrimon, 1999, Bilder et al., 1999, Turenchalk et al., 1999, Kondo, 1998, Woodhouse et al., 1998, Woods et al., 1997, Saha and Sinha, 1996, Caggese et al., 1994, Gateff, 1994, Woodhouse et al., 1994, Bryant, 1993, Karakin et al., 1993, Skaer, 1993, Woods and Bryant, 1992, Szabad et al., 1991, Bryant and Schmidt, 1990, Sokolova and Golubovsky, 1979, Sokolova and Golubovsky, 1979, Sokolova and Golubovsky, 1979, Korochkina et al., 1975)
Name Synonyms
Complementation group 2.1
Lethal (2) giant larva
Lethal giant larva
Lethal(2)giant larvae
Lethal-2-giant larvae
Lethal-giant-larvae
l(2) giant larvae
lethal giant larva
lethal giant larve
lethal(2)-giant larvae
lethal- giant-larvae
lethal-2-giant larvae
lethal-giant-larvae
Secondary FlyBase IDs
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    References (636)