Allele Dmel\ple²

General Information
Symbol	Dmel\ple²	Species	D. melanogaster
Name		FlyBase ID	FBal0013852
Feature type	allele	Associated gene	Dmel\ple

Allele class	loss of function allele
Mutagen	ethyl methanesulfonate
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	loss of function allele (Neckameyer and White, 1993)
Mutagen	ethyl methanesulfonate (Tearle and Nusslein-Volhard, 1987)
Mutations Mapped to the Genome

Type Location Additional Notes References
Associated Sequence Data
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name

UniProtKB/Swiss-Prot
UniProtKB/TrEMBL

Progenitor genotype
Nature of the lesion	Statement Reference Small deletion or point mutation. (Neckameyer and White, 1993)
Cytology
Phenotypic Data
Phenotypic Class
	gravitaxis defective, with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF, ple^FS.Scer\UAS (Riemensperger et al., 2011) gravitaxis defective (with ple^FS.Scer\UAS), with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF (Riemensperger et al., 2011) hypoactive, with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF, ple^FS.Scer\UAS (Riemensperger et al., 2011) hypoactive (with ple^FS.Scer\UAS), with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF (Riemensperger et al., 2011) lethal (with Df(3L)XBB70) (Pendleton et al., 2002) lethal \| embryonic stage (Friggi-Grelin et al., 2003) lethal \| embryonic stage \| recessive (Neckameyer and White, 1993) lethal \| heat sensitive (with ple^ts1) (Pendleton et al., 2002) locomotor rhythm defective, with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF, ple^FS.Scer\UAS (Riemensperger et al., 2011) locomotor rhythm defective (with ple^FS.Scer\UAS), with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF (Riemensperger et al., 2011) phototaxis defective, with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF, ple^FS.Scer\UAS (Riemensperger et al., 2011) phototaxis defective, with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF, ple^Scer\UAS.cFa (Riemensperger et al., 2011) phototaxis defective (with ple^FS.Scer\UAS), with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF (Riemensperger et al., 2011) sleep defective, with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF, ple^FS.Scer\UAS (Riemensperger et al., 2011) sleep defective (with ple^FS.Scer\UAS), with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF (Riemensperger et al., 2011) smell perception defective, with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF, ple^FS.Scer\UAS (Riemensperger et al., 2011) smell perception defective (with ple^FS.Scer\UAS), with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF (Riemensperger et al., 2011) visual behavior defective, with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF, ple^FS.Scer\UAS (Riemensperger et al., 2011) visual behavior defective, with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF, ple^Scer\UAS.cFa (Riemensperger et al., 2011) visual behavior defective (with ple^FS.Scer\UAS), with Scer\GAL4^Ddc.PL, Scer\GAL4^ple.PF (Riemensperger et al., 2011)
Phenotype Manifest In
	embryonic/larval dorsal trunk (Hsouna et al., 2007) embryonic/larval tracheal system (Hsouna et al., 2007)
Detailed Description
	Statement Reference The addition of L-DOPA to food ingested by ple[2] heterozygotes significantly improves food intake. Flies expressing ple[FS.Scer\UAS] in a ple[2] background exhibit reduced walking speed compared to wild-type. Similarly, they cover a shorter distance in a 15 minute period. These flies also respond poorly to mechanical shock (that induces climbing in wild-type flies). Flies expressing ple[FS.Scer\UAS] in a ple[2] background (under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL]) are overall less active and possibly sleep more frequently than controls. In addition, sleeping ple[FS.Scer\UAS]; ple[2] flies respond less frequently to mild and moderate mechanical stimuli, whereas response to strong stimuli is comparable to controls. Flies expressing ple[FS.Scer\UAS] in a ple[2] background (under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL]) show no response to 3-iodotyrosine (whereas controls exhibit decreased activity). In contrast, caffeine is found to decrease resting periods in these flies. Flies expressing ple[FS.Scer\UAS] in a ple[2] background (under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL]) do not show any avoidance to shock-associated odors indicating impaired adversive olfactory learning. Interestingly, these flies show an inverse tendency to choose the shock-associated odor during testing. This effect is significant 2 hours after training. Treatment of flies expressing ple[FS.Scer\UAS] in a ple[2] background (under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL]) with the DA precursor L-DOPA significantly improves their negative geotaxis behavior and olfactory learning. Flies expressing ple[FS.Scer\UAS] in a ple[2] background (under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL]) show no significant response to sucrose. The proboscis extension reflex appears normal in these mutants however, indicating that sugar perception and reflex appetitive reactions are preserved. Quantification of food intake suggests that these flies eat one-third as much as controls, demonstrating that they are markedly hypophagic. The addition of L-DOPA significantly improves this phenotype. Flies expressing ple[FS.Scer\UAS] in a ple[2] background (under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL]) appear to show a lack of phototaxis despite displaying normal electroretinograms and optomotor responses. Flies expressing ple[Scer\UAS.cFa] or ple[FS.Scer\UAS] in a ple[2] background (under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL]) display slightly less accurate visual orientation than wild-type. Similarly, using the detour paradigm, both ple[Scer\UAS.cFa]; ple[2] and ple[FS.Scer\UAS]; ple[2] flies demonstrate a roughly wild-type spatial orientation memory. ple[2] flies expressing either ple[Scer\UAS.cFa] or ple[FS.Scer\UAS] under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL] exhibit normal adult life expectancy. Flies expressing ple[Scer\UAS.cFa] in a ple[2] background (under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL]) demonstrate associative memory similar to wild-type, indicating normal olfactory learning. Flies expressing ple[Scer\UAS.cFa] in a ple[2] background (under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL]) demonstrate partial rescue of ple[2] phototactic behavior. (Riemensperger et al., 2011) Heterozygous embryos have ectopically migrating tracheal branches and tracheal cells. The dorsal trunk is often convoluted. (Hsouna et al., 2007) Heterozygotes have normal locomotor activity at 25^oC. (Pendleton et al., 2002) ple^+t8 rescues mutant embryos to viable adults. Some adults die attempting to free themselves from the pupal case, the greatest lethality occurs during the pharate adult stage. Adults that do eclose are paler than their ple⁺ siblings and they appear weaker in their movements with a tendency to drown in the food medium. Rescued larvae are normal in their response to both light and NaCl. (Neckameyer and White, 1993)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Other
Statement Reference Torsin^KO13, ple²/ple[+] has locomotor behavior defective \| third instar larval stage phenotype (Wakabayashi-Ito et al., 2011)
Phenotype Manifest In
Additional Comments
Genetic Interactions
Statement Reference Double heterozygous Torsin[KO13]/+; ple[2]/+ female larvae show a slight but significant reduction in peristaltic frequency relative to either single heterozygote. (Wakabayashi-Ito et al., 2011)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Rescued by	ple² is rescued by ple^+t11 (Neckameyer et al., 2001) ple² is rescued by ple^Δ2g (Friggi-Grelin et al., 2003) ple² is rescued by Scer\GAL4^C1003/ple^Δ1.Scer\UAS (Friggi-Grelin et al., 2003) ple² is rescued by Scer\GAL4^C1003/ple^Δ2.Scer\UAS (Friggi-Grelin et al., 2003) ple² is rescued by Scer\GAL4^elav-C155/ple^Δ1.Scer\UAS (Friggi-Grelin et al., 2003) ple² is rescued by Scer\GAL4^elav-C155/ple^Δ2.Scer\UAS (Friggi-Grelin et al., 2003) ple² is rescued by Scer\GAL4^ple.PF/ple^FS.Scer\UAS/Scer\GAL4^Ddc.PL (Riemensperger et al., 2011) ple² is rescued by Scer\GAL4^ple.PF/ple^Scer\UAS.cFa (Friggi-Grelin et al., 2003) ple² is rescued by Scer\GAL4^ple.PF/ple^Scer\UAS.cFa/Scer\GAL4^Ddc.PL (Riemensperger et al., 2011) ple² is rescued by Scer\GAL4^ple.PF/ple^Δ2.Scer\UAS (Friggi-Grelin et al., 2003)
Partially rescued by	ple² is partially rescued by ple^+t8 (Neckameyer et al., 2001, Neckameyer and White, 1993)
Not rescued by	ple² is not rescued by ple^{FStrunc.Scer\UAS}/Scer\GAL4^ple.PF/Scer\GAL4^Ddc.PL (Riemensperger et al., 2011) ple² is not rescued by ple^Δ1.Scer\UAS/Scer\GAL4^ple.5' (Friggi-Grelin et al., 2003) ple² is not rescued by ple^Δ1.Scer\UAS/Scer\GAL4^ple.PF (Friggi-Grelin et al., 2003) ple² is not rescued by ple^Δ1g (Friggi-Grelin et al., 2003) ple² is not rescued by Scer\GAL4^ple.5'/ple^Δ2.Scer\UAS (Friggi-Grelin et al., 2003)
Comments	Expression of ple[Scer\UAS.cFa] under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL] fully rescues (to the adult stage) the embryonic lethality normally found in ple[2] mutants. Expression of ple[FStrunc.Scer\UAS] under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL] fails to rescue the embryonic lethality normally found in ple[2] mutants. Expression of ple[FS.Scer\UAS] under the control of both Scer\GAL4[ple.PF] and Scer\GAL4[Ddc.PL] fully rescues (to the adult stage) the embryonic lethality normally found in ple[2] mutants. (Riemensperger et al., 2011) ple[2]-induced embryonic lethality is not rescued by the presence of ple[Δ1g]. The presence of ple[Δ2g] transgene either significantly rescues ple[2] embryonic lethality or full pupal rescue. Few rescued adults are observed with ple[Δ2g], possibly because of a limiting expression level of the transgene at this stage. Expression of ple[Scer\UAS.cFa] in DA cells under the control of Scer\GAL4[ple.PF] leads to full pupal rescue of ple[2]. Expression of ple[Δ1.Scer\UAS] in DA cells under the control of Scer\GAL4[ple.PF] fails to rescue the embryonic lethality seen in ple[2] mutants. Expression of ple[Δ2.Scer\UAS] in DA cells under the control of Scer\GAL4[ple.PF] leads to full pupal rescue of ple[2]. Neuronal expression of ple[Δ1.Scer\UAS] under the control of Scer\GAL4[elav-C155] or Scer\GAL4[C1003] results in significant rescue of ple[2] larvae. Examination of the rescued ple[2] larvae reveals that their mouth hooks and denticle belts are pigmented. Rare adult escapers are observed exhibiting a weak and pale cuticle and much reduced viability. Neuronal expression of ple[Δ2.Scer\UAS] under the control of Scer\GAL4[elav-C155] or Scer\GAL4[C1003] results in significant rescue of ple[2] larvae. Expression of ple[Δ1.Scer\UAS] or ple[Δ2.Scer\UAS] under the control of Scer\GAL4[ple.5'] does not rescue the embryonic lethality seen in ple[2] mutants. (Friggi-Grelin et al., 2003) ple^+t8 rescues mutant embryos to viable adults. (Neckameyer and White, 1993)
Stocks ( 0 )

Notes on Origin
Discoverer

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 6 )
Reported As
Symbol Synonym	ple(2) (Wakabayashi-Ito et al., 2011) ple² (Hsouna et al., 2007, Wakabayashi-Ito et al., 2011) ple^2c1 (Friggi-Grelin et al., 2003) ple^2cl (Friggi-Grelin et al., 2003) ple^7O (Tearle and Nusslein-Volhard, 1987) ple^cl (Riemensperger et al., 2011)
Name Synonym
Secondary FlyBase IDs

References ( 11 )
Research paper	Riemensperger et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(2): 834--839 Behavioral consequences of dopamine deficiency in the Drosophila central nervous system. [FBrf0212779] Wakabayashi-Ito et al., 2011, PLoS ONE 6(10): e26183 dtorsin, the Drosophila Ortholog of the Early-Onset Dystonia TOR1A (DYT1), Plays a Novel Role in Dopamine Metabolism. [FBrf0216459] Hsouna et al., 2007, Dev. Biol. 308(1): 30--43 Drosophila dopamine synthesis pathway genes regulate tracheal morphogenesis. [FBrf0200509] Friggi-Grelin et al., 2003, J. Neurobiol. 54(4): 618--627 Targeted gene expression in Drosophila dopaminergic cells using regulatory sequences from tyrosine hydroxylase. [FBrf0155889] Friggi-Grelin et al., 2003, genesis 35(3): 175--184 Tissue-specific developmental requirements of Drosophila tyrosine hydroxylase isoforms. [FBrf0159367] Pendleton et al., 2002, Behav. Genet. 32(2): 89--94 Effects of tyrosine hydroxylase mutants on locomotor activity in Drosophila: a study in functional genomics. [FBrf0148899] Neckameyer et al., 2001, J. Neurobiol. 47(4): 280--294 Dopamine and sensory tissue development in Drosophila melanogaster. [FBrf0135852] Piedrafita et al., 1994, Insect Biochem. Mol. Biol. 24(6): 581--588 A genetic analysis of aromatic amino acid hydroxylases involvement in DOPA synthesis during Drosophila adult development. [FBrf0074167] Neckameyer and White, 1993, J. Neurogenet. 8(4): 189--199 Drosophila tyrosine hydroxylase is encoded by the pale locus. [FBrf0058913] Budnik and White, 1987, J. Neurogenet. 4: 309--314 Genetic dissection of dopamine and serotonin synthesis in the nervous system of Drosophila melanogaster. [FBrf0046824]
Stock list	Tearle and Nusslein-Volhard, 1987, D. I. S. 66: 209--269 Tubingen mutants and stock list. [FBrf0045941]

Allele Dmel\ple2

Allele Dmel\ple²