D. melanogaster Gene l(2)gl (CG2671-RE)
  Description: CG2671-PE, isoform E (CG2671-PF, isoform F).
Transcript (Including UTRs)
   Position: dm3 chr2L:9,836-21,372 Size: 11,537 Total Exon Count: 9 Strand: -
Coding Region
   Position: dm3 chr2L:11,215-15,648 Size: 4,434 Coding Exon Count: 7 

Page IndexSequence and LinksUniProtKB CommentsMicroarray ExpressionProtein StructureOther Species
GO AnnotationsFlyBase RolesPhenotypesSynonymsIn SitusmRNA Descriptions
Other NamesMethods
Data last updated at UCSC: 2008-10-21

-  Sequence and Links to Tools and Databases
 
Genomic Sequence (chr2L:9,836-21,372)mRNAProtein (1112 aa)
Gene SorterGenome BrowserOther Species FASTATable SchemaFlyBasePubMed
RefSeqUniProtKB

-  Comments and Description Text from UniProtKB
  ID: L2GL_DROME
DESCRIPTION: RecName: Full=Lethal(2) giant larvae protein;
FUNCTION: Essential for the development of polarized epithelia and for cell polarity associated with asymmetric cell division of neuroblasts during development. Isoform p78 has an essential role in control of cell proliferation and differentiation during development and could act as tumor suppressor. Isoform p217 has an accessory function in this respect.
SUBUNIT: May form multimeric complexes.
SUBCELLULAR LOCATION: Cell membrane. Secreted, extracellular space, extracellular matrix. Note=Intercellular matrix.
TISSUE SPECIFICITY: Expressed in the epithelial cells of the digestive tract and in gonads.
DEVELOPMENTAL STAGE: Expressed abundantly in early embryogenesis. Moderate expression is found in larval and adult stages.
SIMILARITY: Belongs to the WD repeat L(2)GL family.
SIMILARITY: Contains 14 WD repeats.

-  Microarray Expression Data
 
Expression ratio colors:

Arbeitman et al. 2002 Life-Cycle Expression Data

     
          
          
     
     
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-  Protein Domain and Structure Information
  InterPro Domains: Graphical view of domain structure
IPR000664 - Lethal2_giant
IPR013905 - Lgl_C
IPR013577 - LLGL2
IPR015943 - WD40/YVTN_repeat-like_dom
IPR001680 - WD40_repeat
IPR017986 - WD40_repeat_dom

Pfam Domains:
PF08596 - Lethal giant larvae(Lgl) like, C-terminal
PF08366 - LLGL2

ModBase Predicted Comparative 3D Structure on P08111
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-  Orthologous Genes in Other Species
  Orthologies between human, mouse, and rat are computed by taking the best BLASTP hit, and filtering out non-syntenic hits. For more distant species reciprocal-best BLASTP hits are used. Note that the absence of an ortholog in the table below may reflect incomplete annotations in the other species rather than a true absence of the orthologous gene.
HumanMouseRatZebrafishC. elegansS. cerevisiae
No orthologNo orthologNo orthologNo orthologNo orthologNo ortholog
      
      
      
      
      

-  Gene Ontology (GO) Annotations with Structured Vocabulary
  Molecular Function:
GO:0017022 myosin binding
GO:0005515 protein binding
GO:0045159 myosin II binding
GO:0005200 structural constituent of cytoskeleton

Biological Process:
GO:0008105 asymmetric protein localization
GO:0045167 asymmetric protein localization involved in cell fate determination
GO:0045175 basal protein localization
GO:0030154 cell differentiation
GO:0008283 cell proliferation
GO:0007391 dorsal closure
GO:0035072 ecdysone-mediated induction of salivary gland cell autophagic cell death
GO:0045197 establishment or maintenance of epithelial cell apical/basal polarity
GO:0045196 establishment or maintenance of neuroblast polarity
GO:0016332 establishment or maintenance of polarity of embryonic epithelium
GO:0016334 establishment or maintenance of polarity of follicular epithelium
GO:0016336 establishment or maintenance of polarity of larval imaginal disc epithelium
GO:0045200 establishment of neuroblast polarity
GO:0045184 establishment of protein localization
GO:0007559 obsolete histolysis
GO:0006886 intracellular protein transport
GO:0001738 morphogenesis of a polarized epithelium
GO:0002009 morphogenesis of an epithelium
GO:0016333 morphogenesis of follicular epithelium
GO:0016335 morphogenesis of larval imaginal disc epithelium
GO:0045746 negative regulation of Notch signaling pathway
GO:0008285 negative regulation of cell proliferation
GO:0007399 nervous system development
GO:0007269 neurotransmitter secretion
GO:0008104 protein localization
GO:0042127 regulation of cell proliferation
GO:0008360 regulation of cell shape
GO:0007423 sensory organ development
GO:0019991 septate junction assembly
GO:0016082 synaptic vesicle priming
GO:0007179 transforming growth factor beta receptor signaling pathway
GO:0045186 zonula adherens assembly

Cellular Component:
GO:0016327 apicolateral plasma membrane
GO:0016323 basolateral plasma membrane
GO:0005938 cell cortex
GO:0005737 cytoplasm
GO:0005856 cytoskeleton
GO:0005829 cytosol
GO:0005578 proteinaceous extracellular matrix
GO:0005614 interstitial matrix
GO:0005886 plasma membrane
GO:0005918 septate junction
GO:0008021 synaptic vesicle


-  FlyBase Biological Roles [Jan. 2006 data]
 
  • Genetic analysis of l(2)gl reveal function is required during embryonic and post-embryonic development to maintain the normal developmental capacity. (Baek and Hanratty, 1997, Cancer Lett. 111(1-2): 233--238)
  • 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. (Woods et al., 1997, Dev. Genet. 20(2): 111--118)
  • l(2)gl plays a critical role at the onset of vitellogenesis and regulates growth of the oocyte, follicle cell migration and organization and germline cell viability. (De Lorenzo et al., 1999, Int. J. Dev. Biol. 43(3): 207--217)
  • 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. (Ohshiro et al., 2000, Nature 408(6812): 593--596)
  • 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. (Peng et al., 2000, Nature 408(6812): 596--600)
  • l(2)gl is required downstream of dpp for the specification of dorsal epidermis. (Arquier et al., 2001, Development 128(12): 2209--2220)
  • l(2)gl promotes cortical localization of mira. (Betschinger et al., 2003, Nature 422(6929): 326--330)
  • l(2)gl acts together with numb in N inhibition and cell fate specification in the adult sensory organ precursor lineage. (Justice et al., 2003, Curr. Biol. 13(9): 778--783)

-  FlyBase Phenotypes of Various Alleles of Gene [Jan. 2006 data]
 
  • 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). (FlyBase)
  • l(2)gl has been cloned and the temporal pattern of RNA expression analyzed. (Mechler et al., 1985, EMBO J. 4: 1551--1557)
  • 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. (Klaembt and Schmidt, 1986, EMBO J. 5(11): 2955--2961)
  • 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. (Kimchie et al., 1989, Cell Diffn Dev. 26: 79--86)
  • Fas3, mys, disco, zip, l(2)gl, N and Egfr mutants show an additive phenotype in combination with Fas1TE89Da. (Elkins et al., 1990, Cell 60: 565--575)
  • 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. (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)
  • awd function is required for brain tumor 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. (Timmons et al., 1993, Dev. Biol. 158(2): 364--379)
  • The highly divergent cis-regulatory elements of Dpsel(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. (Torok et al., 1993, Oncogene 8(6): 1537--1549)
  • 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. (Woodhouse et al., 1994, Cell Growth Diffn 5(2): 151--159)
  • Mutations in l(2)gl cause malignant tumors in the brain and imaginal discs and generate developmental defects in a number of other tissues. Cellular and subcellular localization 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 organization caused by l(2)gl inactivation. Results also suggest the protein is involved in a cytoskeletal-based intercellular communication system directing cell differentiation. (Strand et al., 1994, J. Cell Biol. 127(5): 1345--1360)
  • 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. (Strand et al., 1994, J. Cell Biol. 127(5): 1361--1373)
  • 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. (Merdes et al., 1995, A. Dros. Res. Conf. 36 Suppl.: 16A)
  • 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. (Strand et al., 1995, Europ. Dros. Res. Conf. 14: 277)
  • Mosaic analysis of l(2)gl mutants suggests a role for l(2)gl in cell-cell signaling and interactions. (Agrawal et al., 1995, Dev. Biol. 172(1): 218--229)
  • Interacts genetically with: ecd (data from l(2)glts3) The l(2)gl product is required in vivo in different types of epithelial cells to control their shape during development. (Manfruelli et al., 1996, Development 122(7): 2283--2294)
  • 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-oligomerization of l(2)gl. (Kalmes et al., 1996, J. Cell Sci. 109(6): 1359--1368)
  • 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, organization of the p127 network and its association with other proteins plays critical roles in the control of cell proliferation. (Jakobs et al., 1996, J. molec. Biol. 264(3): 484--496)
  • l(2)gl function is required for proper development during early embryogenesis. (Baek and Hanratty, 1996, D. I. S. 77: 90--92)
  • Interacts genetically with: scrib (data from l(2)glunspecified) (Bilder et al., 2000, Science 289(5476): 113--116)
  • Interacts genetically with: zip Interacts genetically with: numb (data from l(2)glts3) (Ohshiro et al., 2000, Nature 408(6812): 593--596)
  • Interacts genetically with: dpp (data from l(2)glts3) (Arquier et al., 2001, Development 128(12): 2209--2220)
  • Interacts genetically with: crb (data from l(2)gl01433, l(2)gl275, l(2)gl4) Interacts genetically with: sdt (data from l(2)gl275, l(2)gl4) (Tanentzapf and Tepass, 2003, Nature Cell Biol. 5(1): 46--52)
  • Interacts genetically with: N (data from l(2)glunspecified) (Justice et al., 2003, Curr. Biol. 13(9): 778--783)
  • Interacts genetically with: Ras85D (data from l(2)gl4) (Pagliarini and Xu, 2003, Science 302(5648): 1227--1231)
  • Interacts genetically with: CycE (data from l(2)gl23S9, l(2)gl27S3, l(2)glE2S31, l(2)glE6S2) Interacts genetically with: scrib (data from l(2)gl27S3, l(2)gl4, l(2)glE2S31) (Brumby et al., 2004, Genetics 168(1): 227--251)
Allele l(2)gl1:
  • Phenotypic class: lethal | larval stage Phenotypic class: tumorigenic Phenotype manifest in: larval brain Phenotype manifest in: optic lobe Phenotype manifest in: imaginal disc 90% of mutants die as larvae, 2% as pupae. Extreme allele: larval brain and optic lobes become enlarged and disorganized and the imaginal discs are large and clumped. Discs of mutant larvae transplanted into wild-type female abdomens form large contained tumors, whereas transplanted optic primordia from larval brains form invasive neuroblastomas, which grow rapidly, killing the host within 7-14 days. These tumors can be serially cultured in adult abdomens. (Coffman, 2004, Dev. Cell 6(3): 321--327)
  • Phenotype manifest in: larval brain Phenotype manifest in: imaginal disc Phenotype manifest in: larval optic lobe The brain-ventral ganglion complexes of mutant larvae over 5 days old have brain hemispheres which are abnormal in size and morphology. Neuroblastomas of the optic lobes of the brain ganglia and tumourous imaginal discs are present in the mutant larvae. (Nedelcheva et al., 2001, Int. J. Biochem. & Cell Biol. 33(1): 45--51)
Allele l(2)gl2:
  • Like l(2)gl1. RK3. (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
Allele l(2)gl3:
  • l(2)gl3/l(2)gl2 like l(2)gl1/l(2)gl1. (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
Allele l(2)gl4:
  • extreme allele: larval brain and optic lobes become enlarged and disorganized and the imaginal discs large and clumped; when discs of 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. lethal phase: larvae 89.1%, pupae 10.9% (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
  • Clones of mutant tissue grow and metamorphose nearly normally. Results show that the mutation acts autonomously in tissue transplants but non-autonomously in genetic mosaics within an imaginal disc. (Cline, 1976, Genetics 83(3/1): s16)
  • Phenotype manifest in: dorsal mesothoracic disc The wing discs in homozygous larvae form large amorphous tissue masses, which are composed of disorganized sheets and cords of cells. The number of gap junctions is reduced in these discs. (Ryerse and Nagel, 1984, Dev. Biol. 105: 396--403)
  • Hemizygous larvae do not pupariate at 15 or 29oC. (Hanratty, 1984, Roux Arch. dev. Biol. 193(2): 90--97)
  • Phenotypic class: lethal | recessive Phenotype manifest in: egg | germ-line clone No eggs are produced by females containing homozygous germ-line clones. (Szabad et al., 1991, Genetics 127: 525--533)
  • Phenotype manifest in: larval brain The brain ganglia of 4 to 5 day old homozygous larvae appear smaller than those of 4 to 5 day old wild-type larvae, but are normal in shape. The size of the brain increases rapidly from day 6 onwards in homozygous larvae, and after day 8 appears disorganized and forms a large irregular mass. There are fewer replicating nuclei in the brain ganglia of homozygous larvae compared to the brain ganglia of wild-type larvae, up to day 5. 7 day old brain ganglia of homozygous larvae contain more replicating nuclei than the brain ganglia of wild-type 7 day old larvae. The distribution of the replicating nuclei is abnormal. (Roy and Lakhotia, 1991, J. Genet. 70(3): 161--168)
  • Allele class: amorph (Timmons et al., 1993, Dev. Biol. 158(2): 364--379)
  • Phenotypic class: lethal | recessive | pupal stage Phenotype manifest in: larval brain Phenotype manifest in: imaginal disc Phenotype manifest in: lymph gland Homozygous third instar larvae appear transparent and bloated. Pseudopupae are formed, but the animals die in the puparia. The third larval instar brain behaves as an invasive, transplantable lethal neuroblastoma and the imaginal discs develop into benign, transplantable and lethal neoplasms. The cells grow rapidly in a disorganized way when transplanted into adult female hosts. Ultrastructural studies show that the disc cells do not form tight junctions with each other, in contrast to wild-type. The lymph gland is also abnormal; in most cases the single lobes are much enlarged and fused, and contain a large number of fairly undifferentiated round cells which are released into the body cavity and invade the imaginal discs. (Gateff and Schneiderman, 1969, D. I. S. 44: 47--48)
  • Phenotype manifest in: larval brain Larval period is extended. Brains increase in size throughout the extended larval period, and become progressively more abnormal in morphology, failing to remain as an intact tissue. When transplanted into wild-type adult hosts mutant brains can develop into enormous tumors. (Woodhouse et al., 1994, Cell Growth Diffn 5(2): 151--159)
  • Phenotype manifest in: cuticle | somatic clone Phenotype manifest in: wing | anterior/posterior compartment boundary Homozygous clones give rise to tumorous structures with disrupted cuticle patterns. (Mukherjee et al., 1995, Mech. Dev. 51(2-3): 227--234)
  • Phenotype manifest in: wing Phenotype manifest in: dorsal triple row Phenotype manifest in: chemosensory ventral triple row Phenotype manifest in: dorsal double row Phenotype manifest in: ventral double row Phenotype manifest in: crossvein Phenotype manifest in: wing Phenotype manifest in: cuticle l(2)gl4 mutant clones influence the stability of the anterior/posterior compartment boundary in the developing wing. Clones can straddle the boundary, and display loss of pattern due to cell death resulting in scalloping. Surrounding wild type cells show aberrant growth and differentiation. Wing size is reduced and cross veins are lost. Wild type wing can become folded. Some (less than 10%) clones are characterized by unstructured or poorly cuticularized areas, or clustering of mutant cells in vesicles. (Agrawal et al., 1995, Dev. Biol. 172(1): 218--229)
  • Allele class: amorph Can be partially rescued, to produce a small proportion of viable adults, by l(2)glhs.PM. Such rescued female flies show small ovaries and fusion of the germarium to the youngest egg chambers. (Manfruelli et al., 1996, Development 122(7): 2283--2294)
  • Phenotypic class: lethal | larval stage | recessive Phenotypic class: melanotic 'tumor' Phenotype manifest in: Malpighian tubule Phenotype manifest in: imaginal disc Phenotype manifest in: larval brain Phenotype manifest in: cephalopharyngeal skeleton Allele class: amorph Malpighian tubule color: colorless. Larvae exhibit melanotic tumors of the imaginal discs and brain ganglia. l(2)gl4 Abl2 double mutants exhibit a deformed cephalopharyngeal skeleton. (Saha and Sinha, 1996, J. Genet. 75(2): 161--172)
  • Allele class: loss of function The size of the larval imaginal discs of homozygotes is much reduced if they are also homozygous for corto420. (Kodjabachian et al., 1998, EMBO J. 17(4): 1063--1075)
  • Phenotypic class: tumorigenic | recessive The median survival of adult hosts transplanted with hemizygous l(2)gl4/Df(2L)net62 brain fragments is reduced compared to adult hosts transplanted with wild-type brain fragments. Cells from transplanted hemizygous brain fragments are highly metastatic in the wild-type hosts, forming at least one secondary tumor in 87% of cases. Imaginal discs from l(2)gl4/Df(2L)net62 larvae form secondary tumors in 43% of hosts. (Woodhouse et al., 1998, Dev. Genes Evol. 207(8): 542--550)
  • Phenotype manifest in: follicle cell | somatic clone Follicle cell clones homozygous for l(2)gl4 show loss of cell shape and monolayer organization. (Bilder et al., 2000, Science 289(5476): 113--116)
  • Phenotype manifest in: epithelial cell | germ-line clone Phenotype manifest in: neuroblast & embryo | germ-line clone Phenotype manifest in: EL neuron Embryos derived from maternal homozygous germline clones show an early loss of embryonic epithelial apical/basal polarity and neuroblast defects. l(2)gl4 homozygous embryos show a decrease in EL neuron number at stage 17. (Peng et al., 2000, Nature 408(6812): 596--600)
  • Phenotype manifest in: eye disc Phenotype manifest in: wing vein | somatic clone Eye discs are reduced in size. When clones are induced in the wing with the result that the mutant patches affect both wing surfaces wing veins are lost. When the clone only covers one surface of the wing vein differentiation is unaffected. In either case, clones in intervein regions are wild type. (Arquier et al., 2001, Development 128(12): 2209--2220)
  • Phenotype manifest in: follicle cell | somatic clone Clones of homozygous follicle cell undergo extra cell divisions after stage 6 which is accompanied by a loss of apical-basal polarity. The clones frequently lead to the formation of multiple layers of follicle cells. (Deng et al., 2001, Development 128(23): 4737--4746)
  • Misoriented spindles are sometimes observed the neuroblasts of l(2)gl4 embryos. (Petritsch et al., 2003, Dev. Cell 4(2): 273--281)
  • Phenotype manifest in: epithelial cell | somatic clone Phenotype manifest in: tormogen cell | somatic clone Phenotype manifest in: trichogen cell | somatic clone Phenotype manifest in: tormogen cell | supernumerary | somatic clone Phenotype manifest in: trichogen cell | supernumerary | somatic clone Phenotype manifest in: eo neuron | somatic clone Phenotype manifest in: thecogen cell | somatic clone When l(2)gl4w3 clones are made on the notum, large tumors are seen along with disruptions of junctions between epithelial cells. Within these clones, external sensory organs appear malformed containing additional external cells. Most (84%) mutant external sensory organs consist of three sockets and one hair. Clusters of four sockets are also present at a lower (12%) frequency. Both internal cells are seen to be missing most (96%) of the time. (Justice et al., 2003, Curr. Biol. 13(9): 778--783)
  • Phenotype manifest in: follicle cell | ectopic | somatic clone Homozygous mutant clones in the follicle cells leads to multilayered follicle cells. Mutant clones also cause a rough germline boundary and have invasive properties. (Abdelilah-Seyfried et al., 2003, Development 130(9): 1927--1935)
  • Phenotypic class: tumorigenic Phenotype manifest in: larval brain When homozygous mutant larval brain tissue is transplanted into wild-type adult abdomens, the injected tissue proliferates as a primary tumor and invades adjacent tissue, and cells migrate away from the primary tumor to generate widespread metastatic colonies. Homozygous mutant brains are composed of overgrown tissues with loosely adherent cells. (Woodhouse et al., 2003, Proc. Natl. Acad. Sci. USA 100(20): 11463--11468)
  • Phenotype manifest in: border follicle cell Phenotype manifest in: (with l(2)glts3) border follicle cell Delamination and migration of border follicle cells are both accelerated in l(2)gl4 heterozygotes. In l(2)gl4/l(2)glts3 flies the border follicle cells have much reduced polarity. Delamination of these cells is delayed, but their migration is accelerated. (Szafranski and Goode, 2004, Development 131(9): 2023--2036)
Allele l(2)gl110:
  • Phenotypic class: lethal Phenotypic class: tumorigenic Phenotype manifest in: larval brain Phenotype manifest in: optic lobe Phenotype manifest in: imaginal disc 53.6% of mutants die as larvae, 43.4% as pupae. Extreme allele: larval brain and optic lobes become enlarged and disorganized and the imaginal discs are large and clumped. Discs of mutant larvae transplanted into wild-type female abdomens form large contained tumors, whereas transplanted optic primordia from larval brains form invasive neuroblastomas, which grow rapidly, killing the host within 7-14 days. These tumors can be serially cultured in adult abdomens. (Gateff and Mechler, 1989, Crit. Rev. Oncogen. 1: 221--245)
Allele l(2)gl119:
  • Phenotypic class: viable Some l(2)glD150/l(2)gl119 flies are viable. The viability is affected by temperature; the temperature sensitive period is during the third larval stage. (Sokolova and Golubovsky, 1978, D. I. S. 53: 195--196)
  • l(2)glD150/l(2)gl119 transheterozygotes show temperature-sensitive viability. The temperature-sensitive period is in the third larval stage. (Sokolova and Golubovsky, 1979, Genetika, Moscow 15(12): 2175--2182)
Allele l(2)gl138:
  • Phenotypic class: lethal | larval stage Phenotype manifest in: larval brain Phenotype manifest in: imaginal disc 98.0% of mutants die as larvae, 2.0% as pupae. Weak allele: brain and discs are small and rudimentary and grow slowly in transplants. (Sokolova and Golubovsky, 1979, Genetika, Moscow 15(12): 2175--2182)
Allele l(2)gl275:
  • intermediate allele: moderately enlarged brain and discs, which show enhanced growth when transplanted into wild-type females and death of host is delayed. lethal phase: larvae 79.1%, pupae 20.9% (Gateff, 1978, Biol. Rev., Cambr. 53(1): 123--168)
  • Phenotypic class: lethal | recessive Allele class: loss of function (Klaembt and Schmidt, 1986, EMBO J. 5(11): 2955--2961)
  • Phenotypic class: lethal | larval stage | recessive Phenotype manifest in: larval brain Phenotype manifest in: imaginal disc Homozygous larvae die of brain and imaginal disc neoplasms. (Opper et al., 1987, Oncogene 1: 91--96)
  • Phenotypic class: lethal | larval stage | recessive Phenotype manifest in: imaginal disc Tumorous growth of the imaginal discs. (Saha and Sinha, 1996, J. Genet. 75(2): 161--172)
  • Phenotypic class: hyperplastic Neoplastic overgrowth mutant. Imaginal disc tissue shows abnormal distribution of apico-lateral cell junctional markers. (Woods et al., 1997, Dev. Genet. 20(2): 111--118)
Allele l(2)gl309:
  • Phenotypic class: lethal Phenotype manifest in: larval brain Phenotype manifest in: imaginal disc 54.0% of mutants die as larvae, 46.0% as pupae. Intermediate allele: moderately enlarged brain and discs, which show enhanced growth when transplanted into wild-type females and death of host is delayed. (Trunova and Shearn, 2004, A. Dros. Res. Conf. 45: 694A)
Allele l(2)gl314:
  • Phenotypic class: lethal Phenotype manifest in: larval brain Phenotype manifest in: imaginal disc 57.3% of mutants die as larvae, 42.7% as pupae. Weak allele: brain and discs are small and rudimentary and grow slowly in transplants. (Gateff, 1978, Biol. Rev., Cambr. 53(1): 123--168)
Allele l(2)gl334:
  • Phenotypic class: lethal Phenotype manifest in: larval brain Phenotype manifest in: imaginal disc 18.4% of mutants die as larvae, 81.6% as pupae. Weak allele: brain and discs are small and rudimentary and grow slowly in transplants. (Mechler et al., 1985, EMBO J. 4: 1551--1557)
  • Some Df(2L)U558/l(2)gl334 flies are viable. The viability is affected by temperature; the temperature sensitive period is diphasic at 17oC and 29oC. (Sokolova and Golubovsky, 1978, D. I. S. 53: 195--196)
  • l(2)gl558/l(2)gl334 transheterozygotes show temperature-sensitive viability. The temperature-sensitive period is biphasic; the first phase is embryonic and the second phase is in the third larval-prepupal stage. (Sokolova and Golubovsky, 1979, Genetika, Moscow 15(12): 2175--2182)
Allele l(2)gl351:
  • Phenotypic class: lethal Phenotype manifest in: larval brain Phenotype manifest in: imaginal disc 17.5% of mutants die as larvae, 82.5% as pupae. Intermediate allele: moderately enlarged brain and discs, which show enhanced growth when transplanted into wild-type females and death of host is delayed. (Albertson and Doe, 2003, Nature Cell Biol. 5(2): 166--170)
Allele l(2)gl353:
  • Phenotypic class: lethal Phenotype manifest in: larval brain Phenotype manifest in: imaginal disc 19.5% of mutants die as larvae, 80.5% as pupae. Weak allele: brain and discs are small and rudimentary and grow slowly in transplants. (Gateff, 1978, Biol. Rev., Cambr. 53(1): 123--168)
Allele l(2)gl558:
  • l(2)gl558/l(2)gl334 transheterozygotes show temperature-sensitive viability. The temperature-sensitive period is biphasic; the first phase is embryonic and the second phase is in the third larval-prepupal stage. (Mechler et al., 1985, EMBO J. 4: 1551--1557)
  • l(2)gl558/l(2)gl334 transheterozygotes show temperature-sensitive viability. The temperature-sensitive period is biphasic; the first phase is embryonic and the second phase is in the third larval-prepupal stage. (Sokolova and Golubovsky, 1979, Genetika, Moscow 15(12): 2175--2182)
Allele l(2)gl705:
  • Phenotypic class: lethal Phenotype manifest in: larval brain Phenotype manifest in: imaginal disc 89.7% of mutants die as larvae, 10.3% as pupae. Intermediate allele: moderately enlarged brain and discs, which show enhanced growth when transplanted into wild-type females and death of host is delayed. (Sokolova and Golubovsky, 1979, Genetika, Moscow 15(12): 2175--2182)
Allele l(2)gl01433:
  • Phenotype manifest in: epithelial cell Embryos derived from homozygous germline clones (that lack zygotic and maternal l(2)gl function) show defects in epithelial polarity. (Gateff, 1978, Biol. Rev., Cambr. 53(1): 123--168)
  • Phenotype manifest in: epithelial cell Embryos derived from homozygous germline clones (that lack zygotic and maternal l(2)gl function) show defects in epithelial polarity. (Tanentzapf and Tepass, 2003, Nature Cell Biol. 5(1): 46--52)
Allele l(2)gl23S9:
  • Phenotypic class: lethal | larval stage | recessive Lethal phase is during the third larval instar. (Tanentzapf and Tepass, 2003, Nature Cell Biol. 5(1): 46--52)
  • Phenotypic class: lethal | larval stage | recessive Lethal phase is during the third larval instar. (Brumby et al., 2004, Genetics 168(1): 227--251)
Allele l(2)gl27S3:
  • Phenotypic class: lethal | larval stage | recessive Lethal phase is during the third larval instar. (Brumby et al., 2004, Genetics 168(1): 227--251)
  • Phenotypic class: lethal | larval stage | recessive Lethal phase is during the third larval instar. (Brumby et al., 2004, Genetics 168(1): 227--251)
Allele l(2)gl4w3:
  • Phenotype manifest in: follicle cell | somatic clone Follicle cell clones show epithelial defects. (Bilder et al., 2000, Science 289(5476): 113--116)
  • Phenotype manifest in: wing vein | somatic clone When clones are induced in the wing with the result that the mutant patches affect both wing surfaces wing veins are lost. When the clone only covers one surface of the wing vein differentiation is unaffected. In either case, clones in intervein regions are wild type. (Arquier et al., 2001, Development 128(12): 2209--2220)
  • Phenotype manifest in: epithelial cell | somatic clone Phenotype manifest in: tormogen cell | somatic clone Phenotype manifest in: trichogen cell | somatic clone Phenotype manifest in: tormogen cell | supernumerary | somatic clone Phenotype manifest in: trichogen cell | supernumerary | somatic clone Phenotype manifest in: eo neuron | somatic clone Phenotype manifest in: thecogen cell | somatic clone When l(2)gl4w3 clones are made on the notum, large tumors are seen along with disruptions of junctions between epithelial cells. Within these clones, external sensory organs appear malformed containing additional external cells. Most (84%) mutant external sensory organs consist of three sockets and one hair. Clusters of four sockets are also present at a lower (12%) frequency. Both internal cells are seen to be missing most (96%) of the time. (Justice et al., 2003, Curr. Biol. 13(9): 778--783)
Allele l(2)glB:
  • Third instar larvae are bloated and fail to pupate. (Justice et al., 2003, Curr. Biol. 13(9): 778--783)
  • Third instar larvae are bloated and fail to pupate. (Burnet, 1968, D. I. S. 43: 61)
Allele l(2)glD150:
  • Phenotypic class: lethal | larval stage Phenotypic class: tumorigenic Phenotype manifest in: larval brain Phenotype manifest in: optic lobe Phenotype manifest in: imaginal disc 97.3% of mutants die as larvae, 2.7% as pupae. Extreme allele: larval brain and optic lobes become enlarged and disorganized and the imaginal discs are large and clumped. Discs of mutant larvae transplanted into wild-type female abdomens form large contained tumors, whereas transplanted optic primordia from larval brains form invasive neuroblastomas, which grow rapidly, killing the host within 7-14 days. These tumors can be serially cultured in adult abdomens. (Gene Disruption Project members and Exelixis, 2005, computer file)
  • Some l(2)glD150/l(2)gl119 flies are viable. The viability is affected by temperature; the temperature sensitive period is during the third larval stage. (Sokolova and Golubovsky, 1978, D. I. S. 53: 195--196)
  • l(2)glD150/l(2)gl119 transheterozygotes show temperature-sensitive viability. The temperature-sensitive period is in the third larval stage. (Sokolova and Golubovsky, 1979, Genetika, Moscow 15(12): 2175--2182)
Allele l(2)glE2S31:
  • Phenotypic class: lethal | larval stage | recessive Lethal phase is during the third larval instar. (Gene Disruption Project members and Exelixis, 2005, computer file)
  • Phenotypic class: lethal | larval stage | recessive Lethal phase is during the third larval instar. (Brumby et al., 2004, Genetics 168(1): 227--251)
Allele l(2)glE6S2:
  • Phenotypic class: lethal | larval stage | recessive Lethal phase is during the third larval instar. (Brumby et al., 2004, Genetics 168(1): 227--251)
  • Phenotypic class: lethal | larval stage | recessive Lethal phase is during the third larval instar. (Brumby et al., 2004, Genetics 168(1): 227--251)
Allele l(2)glGB652:
  • Phenotypic class: lethal | larval stage Eggs hatch late, there is an extended period of larval growth, and third instar larvae are larger than normal. No larvae survive beyond the third instar stage. (Gene Disruption Project members and Exelixis, 2005, computer file)
  • Phenotypic class: lethal | larval stage Eggs hatch late, there is an extended period of larval growth, and third instar larvae are larger than normal. No larvae survive beyond the third instar stage. (Narachi and Boyd, 1985, Molec. gen. Genet. 199: 500--506)
Allele l(2)glts1:
  • Phenotype manifest in: imaginal disc Phenotype manifest in: dorsal mesothoracic disc Phenotype manifest in: dorsal metathoracic disc Phenotype manifest in: ventral prothoracic disc Phenotype manifest in: ventral metathoracic disc Shows 100% lethality when heterozygous with l(2)gl4 or Df(2L)net62 at 29oC. Does not adversely effect larval viability at either 29oC or 15oC. Hemizygous larvae form a puparium at 29oC, however the majority of larvae fail to evert their imaginal discs and only 3% of larvae differentiate adult cuticle structures. At 15oC, 96% of larvae evert their imaginal discs and 41% of larvae differentiate adult cuticle structures. Imaginal discs are morphologically abnormal in larvae raised at 29oC; approximately 60% of wing discs and 38% of haltere discs are abnormal, and some abnormal first and third leg discs are seen. The proximal region of the wing discs is thickened and has lost its normal folding pattern, while the distal region is normal in shape and has folds. The abnormal regions contains cells arranged in multilayer clumps and chords. Wing discs raised at 29oC and injected into wild-type larvae produce undifferentiated tissue in addition to differentiated tissue and residual material. The undifferentiated tissue is neoplastic when placed in adult hosts at 29oC. (Hanratty, 1984, Roux Arch. dev. Biol. 193(2): 90--97)
  • Homozygous brain hemispheres injected into adult hosts remain normal at 22oC and 29oC. Homozygous larvae raised at 29oC contain both morphologically normal and abnormal wing discs. Discs of both types cultured in adult hosts at 29oC give rise to transplantable neoplastic tissue. When discs raised at 29oC are cultured in adult host at 15oC, the morphologically normal discs maintain their normal morphology, but the morphologically abnormal discs give rise to neoplasms. Homozygous larvae raised at 15oC have morphologically normal wing discs, which maintain their normal morphology when cultured in adult hosts at 15oC, but give rise to neoplasms when cultured in adult hosts at 29oC. Homozygous wing discs raised at 15oC, cut into two fragments and cultured in adult hosts at 15oC become neoplastic. (Hanratty, 1984, Roux Arch. dev. Biol. 193(2): 98--107)
  • Phenotypic class: (with Df(2L)net62) lethal | conditional ts Phenotypic class: (with l(2)glts3) male sterile | conditional ts Phenotype manifest in: (with l(2)glts3) male genitalia Lethality of heterozygotes with Df(2L)net62 when raised at 29oC is complete. (Manfruelli et al., 1996, Development 122(7): 2283--2294)
  • Phenotypic class: lethal | conditional ts Phenotypic class: female sterile | conditional ts Viability of heterozygotes with l(2)glts3 is temperature-dependent, none of the progeny emerged from a cross with l(2)glts2 at 29oC. At 22oC heteroallelic females produce eggs that develop into normal embryos, at 29oC eggs with normal morphology are produced but they fail to hatch into viable larvae. (Baek and Hanratty, 1996, D. I. S. 77: 90--92)
  • Phenotypic class: lethal | maternal effect | conditional ts Phenotype manifest in: dorsal mesothoracic disc | conditional ts Phenotype manifest in: extended germ band embryo | conditional ts | maternal effect The majority of intact wing imaginal discs cultured in vivo at 15oC remain morphologically normal but show no significant growth. The majority of intact discs grown at 29oC become abnormal and show signs of growth. Culture of amputated discs at 15oC generates abnormal discs. Gamma irradiated discs (whole or amputated) become abnormal after in vivo culture at 15oC but show no significant growth. l(2)glts1/l(2)glts3 heteroallelic females exhibit normal morphology of ovaries and developing egg chambers. Heterozygous females lay normal eggs at 22oC and 29oC. At 29oC the eggs fail to develop into viable larvae. None of the unhatched eggs show differentiated larval cuticles. Embryos stop developing at the stage of germ band formation, none of the embryos exhibit dorsal closure and differentiation of the larval hypodermis. None of the embryos have developed a well defined CNS. (Baek and Hanratty, 1997, Cancer Lett. 111(1-2): 233--238)
Allele l(2)glts2:
  • Phenotype manifest in: imaginal disc Shows 100% lethality when heterozygous with l(2)gl4 or Df(2L)net62 at 29oC. Larval viability is significantly reduced at 29oC or 15oC. Hemizygous larvae form a puparium at 29oC, however 90% of larvae fail to evert their imaginal discs and only 7% of larvae differentiate adult cuticle structures. At 15oC, 55% of larvae evert their imaginal discs and 14% of larvae differentiate adult cuticle structures. (Hanratty, 1984, Roux Arch. dev. Biol. 193(2): 90--97)
  • Phenotypic class: (with Df(2L)net62) lethal | conditional ts Phenotypic class: (with l(2)glts3) male sterile | conditional ts Phenotype manifest in: (with l(2)glts3) male genitalia Lethality of heterozygotes with Df(2L)net62 when raised at 29oC is complete. (Manfruelli et al., 1996, Development 122(7): 2283--2294)
  • Phenotypic class: lethal | conditional ts Phenotypic class: female sterile | conditional ts Viability of heterozygotes with l(2)glts3 is temperature-dependent, none of the progeny emerged from a cross with l(2)glts1 at 29oC. At 22oC heteroallelic females produce eggs that develop into normal embryos, at 29oC eggs with normal morphology are produced but they fail to hatch into viable larvae. (Baek and Hanratty, 1996, D. I. S. 77: 90--92)
  • Phenotypic class: lethal | maternal effect | conditional ts Phenotype manifest in: dorsal mesothoracic disc | conditional ts Phenotype manifest in: extended germ band embryo | conditional ts | maternal effect The majority of intact wing imaginal discs cultured in vivo at 15oC remain morphologically normal but show no significant growth. The majority of intact discs grown at 29oC become abnormal and show signs of growth. Culture of amputated discs at 15oC generates abnormal discs. Gamma irradiated discs (whole or amputated) become abnormal after in vivo culture at 15oC but show no significant growth. l(2)glts2/l(2)glts3 heteroallelic females exhibit normal morphology of ovaries and developing egg chambers. Heterozygous females lay normal eggs at 22oC and 29oC. At 29oC the eggs fail to develop into viable larvae. None of the unhatched eggs show differentiated larval cuticles. Embryos stop developing at the stage of germ band formation, none of the embryos exhibit dorsal closure and differentiation of the larval hypodermis. None of the embryos have developed a well defined CNS. (Baek and Hanratty, 1997, Cancer Lett. 111(1-2): 233--238)
Allele l(2)glts3:
  • Phenotypic class: (with Df(2L)net62) lethal | conditional ts Phenotypic class: (with l(2)glts1) male sterile | conditional ts Phenotypic class: (with l(2)glts2) male sterile | conditional ts Phenotypic class: female sterile | recessive | conditional ts Phenotype manifest in: (with l(2)glts1) male genitalia Phenotype manifest in: (with l(2)glts2) male genitalia Phenotype manifest in: extended germ band embryo embryo Phenotype manifest in: embryonic head Phenotype manifest in: larval midgut & embryo Phenotype manifest in: egg chamber Phenotype manifest in: follicle cell Phenotype manifest in: germarium Allele class: hypomorph Lethality of heterozygotes with Df(2L)net62 when raised at 29oC is complete. Homozygous mutant larvae produced at 29oC pupariate on a normal timetable and die during the early pupal stages. Male homozygotes and heterozygotes with l(2)glts1 or l(2)glts2 show male sterility when raised at 27oC. Genitalia are incompletely rotated, disorganized and protruding. Mutant embryos from mutant mothers show developmental block at germ band shortening. Head involution fails but segmentation progresses normally. Cell shape changes at the heading edge of the dorsal epidermis fail to occur, reminiscent of zip and scb mutant phenotypes. Midgut is hypertrophied and the yolk remains incompletely digested. The visceral mesoderm is apparently normal. Midgut cells arrest their differentiation early. Causes a temperature sensitive reduction in female fertility. Oogenesis blocks at stage 7-8 in females raised at 29oC for 6 days. Blocked egg chambers show a multilayered accumulation of probably follicular cells at the anterior and posterior tips. These cells become internalized into the space normally occupied by the ooplasm. The germarium becomes fused with the youngest egg chambers. (Manfruelli et al., 1996, Development 122(7): 2283--2294)
  • Phenotypic class: lethal | conditional ts Phenotypic class: female sterile | conditional ts Viability of heterozygotes with l(2)glts1 and l(2)glts2 is temperature-dependent. At 22oC heteroallelic females produce eggs that develop into normal embryos, at 29oC eggs with normal morphology are produced but they fail to hatch into viable larvae. (Baek and Hanratty, 1996, D. I. S. 77: 90--92)
  • Phenotypic class: lethal | maternal effect | conditional ts Phenotype manifest in: dorsal mesothoracic disc | conditional ts Phenotype manifest in: extended germ band embryo | conditional ts | maternal effect The majority of intact wing imaginal discs cultured in vivo at 15oC remain morphologically normal but show no significant growth. The majority of intact discs grown at 29oC become abnormal and show signs of growth. Culture of amputated discs at 15oC generates abnormal discs. Gamma irradiated discs (whole or amputated) become abnormal after in vivo culture at 15oC but show no significant growth. l(2)glts1/l(2)glts3 and l(2)glts2/l(2)glts3 heteroallelic females exhibit normal morphology of ovaries and developing egg chambers. Heterozygous females lay normal eggs at 22oC and 29oC. At 29oC the eggs fail to develop into viable larvae. None of the unhatched eggs show differentiated larval cuticles. Embryos stop developing at the stage of germ band formation, none of the embryos exhibit dorsal closure and differentiation of the larval hypodermis. None of the embryos have developed a well defined CNS. (Baek and Hanratty, 1997, Cancer Lett. 111(1-2): 233--238)
  • Phenotype manifest in: egg chamber Phenotype manifest in: ovary Ovaries of l(2)glts3 females shifted to 29oC immediately after hatching and reared at 29oC for 2 days are severely reduced in size, due to a lack of late stage egg chambers. (De Lorenzo et al., 1999, Int. J. Dev. Biol. 43(3): 207--217)
  • Phenotype manifest in: external sensory organ Phenotype manifest in: socket Phenotype manifest in: tormogen cell Phenotype manifest in: eo neuron Exposure to 29oC during external sensory organ development causes transformation of inner cells to outer cells causing double socket and hair phenotypes. (Ohshiro et al., 2000, Nature 408(6812): 593--596)
  • Phenotype manifest in: embryonic epidermis Phenotype manifest in: embryonic ventral epidermis Phenotype manifest in: embryonic leading edge cell Phenotype manifest in: cephalopharyngeal skeleton Phenotype manifest in: filzkorper Phenotype manifest in: denticle belt The polarity of the dorsal leading edge cells of the embryonic epidermis is disrupted, though other ectodermal cells are normal. Embryos reared at 29oC show failures of dorsal closure and failure of head involution. Cephalopharyngeal skeleton and filzkorper can be absent or not externalized, respectively. 58% of cuticles show a dorsal hole. Denticle belts are extended, reaching up to 60% of lateral extension in some mutant individuals. Rare embryos show the phenotype of partly lacking ventral epidermis. Less than 8% of cuticles show patches of denticle belt fusion or naked cuticle, indicating that on the whole wg and hh signaling are unaffected. (Arquier et al., 2001, Development 128(12): 2209--2220)
  • Phenotype manifest in: (with l(2)gl4) border follicle cell In l(2)gl4/l(2)glts3 flies the border follicle cells have much reduced polarity. Delamination of these cells is delayed, but their migration is accelerated. (Szafranski and Goode, 2004, Development 131(9): 2023--2036)
Allele l(2)glunspecified:
  • Phenotypic class: lethal | recessive Phenotype manifest in: prothoracic gland The prothoracic gland cells of homozygous 140 hour old third instar larvae are smaller than normal. Glycogen deposits, secretory droplets and lysosomes, which are normally seen in wild-type cells, are absent. The volume of smooth endoplasmic reticulum (ER) is reduced to approximately 1% of that seen in wild-type, and the volume of rough ER is reduced to about 20% of the normal value. The corpus allatum appears fairly normal, although secretory deposits are not seen. (Aggarwal and King, 1969, J. Morph. 129: 171--200)
  • Phenotype manifest in: larval corpus allatum Phenotype manifest in: prothoracic gland Phenotype manifest in: larval salivary gland & embryo Larvae homozygous for a l(2)glunspecified allele that is lethal during the larval stage have underdeveloped prothoracal glands, although the corpora allata are normal. Salivary glands are 50% of normal size and polyteny is reduced. Larvae homozygous for a l(2)glunspecified allele that is lethal during the prepupal stage have underdeveloped prothoracal glands and reduced corpora allata. Salivary glands are 80% of normal size and polyteny is reduced. (Korochkina et al., 1975, Genetika, Moscow 11: 57--65)
  • Phenotype manifest in: adherens junction Embryos lacking maternal and zygotic l(2)gl function show defects in apicobasal polarity during mid-embryogenesis and disruption of adherens junctions. (Bilder et al., 2000, Science 289(5476): 113--116)
  • Neuroblasts in homozygous stage 15 embryos (lacking zygotic l(2)gl function) show defects in division; 57% undergo normal division, 31% undergo a symmetrical division and 12% undergo an inverted asymmetrical division. (Albertson and Doe, 2003, Nature Cell Biol. 5(2): 166--170)
  • An ultrastructurally normal junctional complex composed of a zonula adherens and a septate junction is seen in stage 17 embryos which lack maternal and zygotic l(2)gl function. (Tanentzapf and Tepass, 2003, Nature Cell Biol. 5(1): 46--52)
  • Phenotype manifest in: tormogen cell | somatic clone Phenotype manifest in: trichogen cell | somatic clone Phenotype manifest in: tormogen cell | supernumerary | somatic clone Phenotype manifest in: trichogen cell | supernumerary | somatic clone Phenotype manifest in: eo neuron | somatic clone Phenotype manifest in: thecogen cell | somatic clone (Justice et al., 2003, Curr. Biol. 13(9): 778--783)
Allele l(2)gl+t23:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Jacob et al., 1987, Cell 50: 215--225)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Gateff and Mechler, 1989, Crit. Rev. Oncogen. 1: 221--245)
Allele l(2)gl+t13.1:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Jacob et al., 1987, Cell 50: 215--225)
  • Phenotypic class: wild-type Mode of assay: In transgenic Drosophila (intraspecific) (Opper et al., 1987, Oncogene 1: 91--96)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Gateff and Mechler, 1989, Crit. Rev. Oncogen. 1: 221--245)
  • Phenotypic class: wild-type Mode of assay: In transgenic Drosophila (intraspecific) Suppresses the tumor phenotype of l(2)gl mutations. (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)
  • Phenotypic class: wild-type (Saha and Sinha, 1996, J. Genet. 75(2): 161--172)
Allele l(2)gl+tMa:
  • Phenotypic class: wild-type Mode of assay: In transgenic Drosophila (intraspecific) Suppresses the tumor phenotype of l(2)gl mutations. (Saha and Sinha, 1996, J. Genet. 75(2): 161--172)
  • Phenotypic class: wild-type Mode of assay: In transgenic Drosophila (intraspecific) Suppresses the tumor phenotype of l(2)gl mutations. (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)
Allele l(2)gl+tMb:
  • Phenotypic class: wild-type Mode of assay: In transgenic Drosophila (intraspecific) Suppresses the tumor phenotype of l(2)gl mutations. (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)
  • Phenotypic class: wild-type Mode of assay: In transgenic Drosophila (intraspecific) Suppresses the tumor phenotype of l(2)gl mutations. (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)
Allele l(2)gl+tMd:
  • Phenotypic class: lethal | partially | pharate adult stage Phenotype manifest in: pupa Mode of assay: In transgenic Drosophila (intraspecific) Tumor phenotype of l(2)gl deficient animals is rescued, but viability is reduced, with about 60% of animals dying as pharate adults, unable to emerge from the pupal case. (Jacob et al., 1987, Cell 50: 215--225)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Gateff and Mechler, 1989, Crit. Rev. Oncogen. 1: 221--245)
  • Phenotypic class: wild-type Mode of assay: In transgenic Drosophila (intraspecific) Suppresses the tumor phenotype of l(2)gl mutations. (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)
Allele l(2)gl3A.ScerUAS:
  • Mode of assay: In transgenic Drosophila (intraspecific) Expression of l(2)gl3A.ScerUAS by ScerGAL4mat.&agr;Tub67C.T:HsimVP16 does not affect spindle orientation of embryonic neuroblasts. (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)
  • Mode of assay: In transgenic Drosophila (intraspecific) Expression of l(2)gl3A.ScerUAS by ScerGAL4mat.&agr;Tub67C.T:HsimVP16 does not affect spindle orientation of embryonic neuroblasts. (Betschinger et al., 2003, Nature 422(6929): 326--330)
Allele l(2)gl3A.ScerUAS.T:HsapMYC:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Barros et al., 2003, Dev. Cell 5(6): 829--840)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2003, Nature 422(6929): 326--330)
Allele l(2)glC+&bgr;.ScerUAS.T:HsapMYC:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2003, Nature 422(6929): 326--330)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
Allele l(2)glC.ScerUAS.T:HsapMYC:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
Allele l(2)glF311:
  • Phenotypic class: female sterile | conditional ts Phenotypic class: wild-type Phenotype manifest in: egg chamber | conditional ts Phenotype manifest in: ovary | conditional ts Phenotype manifest in: nurse cell | conditional ts Phenotype manifest in: stage S9 oocyte | conditional ts Phenotype manifest in: follicle cell | conditional ts Phenotype manifest in: border follicle cell | conditional ts Phenotype manifest in: oocyte | conditional ts Phenotype manifest in: oocyte & cytoskeleton | conditional ts Phenotype manifest in: nurse cell & cytoskeleton | conditional ts Mode of assay: In transgenic Drosophila (intraspecific) Fully rescues the l(2)gl4 phenotype when raised at the permissive temperature of 22oC. However, drastic effects on the development of these flies are seen at the restrictive temperature of 29oC, in particular the fertility of females is strongly reduced. The fertility of males is not affected. Newly hatched females raised at 22oC and then immediately shifted to 29oC are unable to produce eggs. Females maintained at 22oC for 2-3 days after eclosion and then shifted to 29oC show a nearly normal rate of egg laying during the first day at 29oC, with the rate of egg laying gradually decreasing to about 5% of the control rate after 3 days at 29oC. Eggs laid by females at 29oC have normal morphology and can be fertilized, although their development is blocked when the eggs are maintained at this temperature. Eggs laid for 2 hours by females maintained at 29oC and then incubated at 22oC show normal development. The morphology of ovaries of females carrying l(2)glF311 in a l(2)gl4 background and reared at 22oC is indistinguishable from wild type. Ovaries of females carrying l(2)glF311 in a l(2)gl4 background, shifted to 29oC immediately after hatching and reared at 29oC for 2 days are severely reduced in size, due to a lack of late stage egg chambers. Ovaries of 5 day old females carrying l(2)glF311 in a l(2)gl4 background, shifted to 29oC immediately after hatching and reared at 29oC for 3 days show a marked depletion of egg chambers beyond stage 9. A few ovarioles which have a nearly mature egg are seen, but in these cases the egg is found to follow a degenerating egg chamber. In all other ovarioles the most developed egg chambers show degenerating characteristics (a reduction in size, abnormally arranged nurse cells and a disorganized cytoskeleton) while early stage egg chambers appear normal. The most advanced egg chambers reach stage 9 with a large proportion of the follicle cells being located over the oocyte. The stage 9 egg chamber is smaller than wild type. The layer of follicle cells over the oocyte form an antero-posteior gradient of increasingly elongated cells with an irregular shape and irregularly spaced nuclei. No sharp transition between columnar cells and squamous cells can be detected. The columnar follicle cells over the posterior end of the egg chamber can become extremely elongated. The border follicle cells reach their final destination, as seen by their lateral alignment, however they remain located within the layer of nurse cells bordering the oocyte and are never seen apposed to the anterior border of the oocyte. The nuclei of the most distally located egg chambers in more than half of the ovarioles are very condensed and fragmented, and the cytoskeleton of the nurse cells and oocyte is disrupted in the most advanced degenerating egg chambers. Apoptotic cell death is seen in the degenerating egg chambers. (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
  • Phenotypic class: female sterile | conditional ts Phenotypic class: wild-type Phenotype manifest in: egg chamber | conditional ts Phenotype manifest in: ovary | conditional ts Phenotype manifest in: nurse cell | conditional ts Phenotype manifest in: stage S9 oocyte | conditional ts Phenotype manifest in: follicle cell | conditional ts Phenotype manifest in: border follicle cell | conditional ts Phenotype manifest in: oocyte | conditional ts Phenotype manifest in: oocyte & cytoskeleton | conditional ts Phenotype manifest in: nurse cell & cytoskeleton | conditional ts Mode of assay: In transgenic Drosophila (intraspecific) Fully rescues the l(2)gl4 phenotype when raised at the permissive temperature of 22oC. However, drastic effects on the development of these flies are seen at the restrictive temperature of 29oC, in particular the fertility of females is strongly reduced. The fertility of males is not affected. Newly hatched females raised at 22oC and then immediately shifted to 29oC are unable to produce eggs. Females maintained at 22oC for 2-3 days after eclosion and then shifted to 29oC show a nearly normal rate of egg laying during the first day at 29oC, with the rate of egg laying gradually decreasing to about 5% of the control rate after 3 days at 29oC. Eggs laid by females at 29oC have normal morphology and can be fertilized, although their development is blocked when the eggs are maintained at this temperature. Eggs laid for 2 hours by females maintained at 29oC and then incubated at 22oC show normal development. The morphology of ovaries of females carrying l(2)glF311 in a l(2)gl4 background and reared at 22oC is indistinguishable from wild type. Ovaries of females carrying l(2)glF311 in a l(2)gl4 background, shifted to 29oC immediately after hatching and reared at 29oC for 2 days are severely reduced in size, due to a lack of late stage egg chambers. Ovaries of 5 day old females carrying l(2)glF311 in a l(2)gl4 background, shifted to 29oC immediately after hatching and reared at 29oC for 3 days show a marked depletion of egg chambers beyond stage 9. A few ovarioles which have a nearly mature egg are seen, but in these cases the egg is found to follow a degenerating egg chamber. In all other ovarioles the most developed egg chambers show degenerating characteristics (a reduction in size, abnormally arranged nurse cells and a disorganized cytoskeleton) while early stage egg chambers appear normal. The most advanced egg chambers reach stage 9 with a large proportion of the follicle cells being located over the oocyte. The stage 9 egg chamber is smaller than wild type. The layer of follicle cells over the oocyte form an antero-posteior gradient of increasingly elongated cells with an irregular shape and irregularly spaced nuclei. No sharp transition between columnar cells and squamous cells can be detected. The columnar follicle cells over the posterior end of the egg chamber can become extremely elongated. The border follicle cells reach their final destination, as seen by their lateral alignment, however they remain located within the layer of nurse cells bordering the oocyte and are never seen apposed to the anterior border of the oocyte. The nuclei of the most distally located egg chambers in more than half of the ovarioles are very condensed and fragmented, and the cytoskeleton of the nurse cells and oocyte is disrupted in the most advanced degenerating egg chambers. Apoptotic cell death is seen in the degenerating egg chambers. (De Lorenzo et al., 1999, Int. J. Dev. Biol. 43(3): 207--217)
Allele l(2)glhs.PM:
  • Mode of assay: In transgenic Drosophila (intraspecific) (De Lorenzo et al., 1999, Int. J. Dev. Biol. 43(3): 207--217)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Manfruelli et al., 1996, Development 122(7): 2283--2294)
Allele l(2)glM.ScerUAS.T:HsapMYC:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Manfruelli et al., 1996, Development 122(7): 2283--2294)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
Allele l(2)glMC.ScerUAS.T:HsapMYC:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
Allele l(2)glN.ScerUAS.T:HsapMYC:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
Allele l(2)glScerUAS.cBa:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2003, Nature 422(6929): 326--330)
Allele l(2)glScerUAS.cMa:
  • Mode of assay: In transgenic Drosophila (intraspecific) When expressed under the influence of ScerGAL469B, l(2)glScerUAS.cMa can rescue the lethality of l(2)gl4 mutants, though some of rescued flies show sterility. (Betschinger et al., 2005, Curr. Biol. 15(3): 276--282)
  • Mode of assay: In transgenic Drosophila (intraspecific) When expressed under the influence of ScerGAL469B, l(2)glScerUAS.cMa can rescue the lethality of l(2)gl4 mutants, though some of rescued flies show sterility. (Manfruelli et al., 1996, Development 122(7): 2283--2294)
Allele l(2)glScerUAS.cOa:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Arquier et al., 2001, Development 128(12): 2209--2220)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Ohshiro et al., 2000, Nature 408(6812): 593--596)
Allele l(2)glScerUAS.T:HsapMYC:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Ohshiro et al., 2000, Nature 408(6812): 593--596)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Betschinger et al., 2003, Nature 422(6929): 326--330)
Allele l(2)glt16:
  • Mode of assay: In transgenic Drosophila (intraspecific) Fails to suppress the tumor phenotype of l(2)gl mutations. (Betschinger et al., 2003, Nature 422(6929): 326--330)
  • Mode of assay: In transgenic Drosophila (intraspecific) Fails to suppress the tumor phenotype of l(2)gl mutations. (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)
Allele l(2)glt21:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Jacob et al., 1987, Cell 50: 215--225)
  • Mode of assay: In transgenic Drosophila (intraspecific) Fails to suppress the tumor phenotype of l(2)gl mutations. (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)
Allele l(2)glt22:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Jacob et al., 1987, Cell 50: 215--225)
Allele l(2)glt25:
  • Mode of assay: In transgenic Drosophila (intraspecific) (Jacob et al., 1987, Cell 50: 215--225)
  • Mode of assay: In transgenic Drosophila (intraspecific) (Gateff and Mechler, 1989, Crit. Rev. Oncogen. 1: 221--245)
  • Mode of assay: In transgenic Drosophila (intraspecific) Fails to suppress the tumor phenotype of l(2)gl mutations. (Merz et al., 1990, Environ. Hlth Perspect. 88: 163--167)

-  FlyBase Synonyms for Gene and Its Alleles [Jan. 2006 data]
  Symbol: l(2)gl Name: lethal (2) giant larvae BDGP: CG2671-RE FlyBase: FBgn0002121
Synonyms: 119, 138, 23S9, 275, 27S3, 309, 314, 334, 351, 353, 558, 705, CG2671, Complementation group 2.1, D-LGL, D150, E2S31, E6S2, L(2)GL, LGL, Lethal Giant Larvae, Lgl, dlgl, l(2), l(2) giant larva, l(2)01433, l(2)PM13, l(2)R2, l(2)giant larvae, l(2)gl+, l(2)gl+t13.1, l(2)gl+t23, l(2)gl+tMa, l(2)gl+tMb, l(2)gl+tMd, l(2)gl01433, l(2)gl1, l(2)gl110, l(2)gl119, l(2)gl119, l(2)gl138, l(2)gl150, l(2)gl2, l(2)gl23S9, l(2)gl25, l(2)gl275, l(2)gl27S3, l(2)gl3, l(2)gl309, l(2)gl314, l(2)gl334, l(2)gl334, l(2)gl351, l(2)gl353, l(2)gl3A.ScerUAS, l(2)gl3A.ScerUAS.T:HsapMYC, l(2)gl4, l(2)gl432, l(2)gl4w3, l(2)gl52, l(2)gl558, l(2)gl6, l(2)gl705, l(2)glB, l(2)glC+beta.ScerUAS.T:HsapMYC, l(2)glC.ScerUAS.T:HsapMYC, l(2)glD150, l(2)glDV110, l(2)glDV275, l(2)glDV275, l(2)glE2S31, l(2)glE432, l(2)glE6S2, l(2)glF311, l(2)glGB652, l(2)glI50, l(2)glII0, l(2)glKG05323, l(2)glM.ScerUAS.T:HsapMYC, l(2)glM25, l(2)glMC.ScerUAS.T:HsapMYC, l(2)glN.ScerUAS.T:HsapMYC, l(2)glPM13, l(2)glR2, l(2)glScerUAS.T:HsapMYC, l(2)glScerUAS.cBa, l(2)glScerUAS.cMa, l(2)glScerUAS.cOa, l(2)glU138, l(2)glU314, l(2)glU334, l(2)glU334, l(2)glU353, l(2)glUAS.cMa, l(2)glc07179, l(2)gle04278, l(2)glf02181, l(2)glhs.PM, l(2)glt16, l(2)glt21, l(2)glt22, l(2)glt24, l(2)glt25, l(2)glts1, l(2)glts2, l(2)glts3, l(2)glts3, l(2)glunspecified, l-gl, l2gl, l2gl4, lethal giant larva, lethal giant larvae, lethal giant larve, lethal(2)giant larvae, lethal-giant-larvae, lgl1, lgl334, lgl4, lgl4w3, lgl: lethal-giant-larvae, lglDV275, lgldv275, lglts3, p127, p127l(2)gl, unspecified

-  BDGP Expression In Situ Images
  In situ images for BDGP gene CG2671 (l(2)gl, FlyBase gene FBgn0002121) are available from the BDGP Gene Expression project.
Number of images: 4
Number of body parts: 1
EST: PC00404
DGC Plate: pcr_5
Plate position: 20

-  Descriptions from all associated GenBank mRNAs
  BT100228 - Drosophila melanogaster RE74125 full insert cDNA.
X05426 - Drosophila mRNA for lethal(2) giant larvae protein l(2)gl.
BT100129 - Drosophila melanogaster RE31203 full insert cDNA.
AY051654 - Drosophila melanogaster LD06034 full length cDNA.
FJ634679 - Synthetic construct Drosophila melanogaster clone BS02819 encodes l(2)gl-RA.
FJ629585 - Synthetic construct Drosophila melanogaster clone BO03719 encodes l(2)gl-RA.
JV213333 - TSA: Drosophila melanogaster isotig05180.Dmov mRNA sequence.
JV228107 - TSA: Drosophila melanogaster isotig10413.Dmte mRNA sequence.
KX806240 - Synthetic construct clone BS20569 l(2)gl-PB (l(2)gl) gene, complete cds.
KX794904 - Synthetic construct clone BO27569 l(2)gl-RB (l(2)gl) gene, partial cds.

-  Other Names for This Gene
  UCSC ID: CG2671-RE
RefSeq Accession: NM_164351
Protein: P08111 (aka L2GL_DROME)

-  FlyBase Genes Methods, Credits, and Data Use Restrictions
  Click here for details on how this gene model was made and data restrictions if any.