Allele Dmel\lark¹

General Information
Symbol	Dmel\lark¹	Species	D. melanogaster
Name		FlyBase ID	FBal0035182
Feature type	allele	Associated gene	Dmel\lark

Allele class
Mutagen	Delta2-3, P-element activity
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class
Mutagen	Delta2-3 (Newby and Jackson, 1993) P-element activity (Newby and Jackson, 1996)
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 P{lacW} insertion within a 5' untranslated exon. (Newby and Jackson, 1996)
Caused by insertion	P{lacW}lark¹ (Newby and Jackson, 1993, McNeil et al., 1999, Newby and Jackson, 1996)
Cytology
Phenotypic Data
Phenotypic Class
	behavior defective \| dominant (Newby and Jackson, 1993) eclosion defective \| dominant (Sundram et al., 2012) female fertile, with lark^{cMa.T:Ivir\HA1} (McNeil et al., 2001) female fertile, with lark^{RRM1.RNP2.T:Ivir\HA1} (McNeil et al., 2001) female sterile, with lark^{RRM1+RTZF.T:Ivir\HA1} (McNeil et al., 2001) female sterile, with lark^{RRM2.RNP2.T:Ivir\HA1} (McNeil et al., 2001) female sterile, with lark^{RRM2+RTZF.T:Ivir\HA1} (McNeil et al., 2001) flight defective, with lark^{RRM2.RNP2.T:Ivir\HA1} (McNeil et al., 2001) flightless, with lark^{RRM1+RTZF.T:Ivir\HA1} (McNeil et al., 2001) flightless, with lark^{RRM2+RTZF.T:Ivir\HA1} (McNeil et al., 2001) flightless, with lark^{ZF.T:Ivir\HA1} (McNeil et al., 2001) lethal (Sundram et al., 2012) lethal \| embryonic stage \| recessive (Newby and Jackson, 1993) lethal \| recessive (McNeil et al., 2001) locomotor rhythm defective \| heat sensitive, with Scer\GAL4^dimm-929, lark^{dsRNA.Scer\UAS.cSb} (Sundram et al., 2012) locomotor rhythm defective \| heat sensitive, with Scer\GAL4^elav-C155, lark^{dsRNA.Scer\UAS.cSb} (Sundram et al., 2012) locomotor rhythm defective \| heat sensitive, with Scer\GAL4^Mai179, lark^{dsRNA.Scer\UAS.cSb} (Sundram et al., 2012) locomotor rhythm defective \| heat sensitive, with Scer\GAL4^nrv2.PS, lark^{dsRNA.Scer\UAS.cSb} (Sundram et al., 2012) locomotor rhythm defective \| heat sensitive, with Scer\GAL4^P2.4.Pdf, lark^{dsRNA.Scer\UAS.cSb} (Sundram et al., 2012) locomotor rhythm defective \| heat sensitive, with Scer\GAL4^tim.Scer\UAS, lark^{dsRNA.Scer\UAS.cSb} (Sundram et al., 2012) male fertile, with lark^{cMa.T:Ivir\HA1} (McNeil et al., 2001) male fertile, with lark^{RRM1.RNP2.T:Ivir\HA1} (McNeil et al., 2001) male fertile, with lark^{RRM1+RTZF.T:Ivir\HA1} (McNeil et al., 2001) male fertile, with lark^{RRM2.RNP2.T:Ivir\HA1} (McNeil et al., 2001) male fertile, with lark^{RRM2+RTZF.T:Ivir\HA1} (McNeil et al., 2001) visible, with lark^{RRM2.RNP2.T:Ivir\HA1} (McNeil et al., 2001) visible, with lark^{RRM2+RTZF.T:Ivir\HA1} (McNeil et al., 2001) visible, with lark^{ZF.T:Ivir\HA1} (McNeil et al., 2001)
Phenotype Manifest In
	embryo \| germline clone (McNeil et al., 1999) posterior scutellar bristle, with lark^{ZF.T:Ivir\HA1} (McNeil et al., 2001) scutellar bristle, with lark^{RRM2.RNP2.T:Ivir\HA1} (McNeil et al., 2001) scutellar bristle, with lark^{RRM2+RTZF.T:Ivir\HA1} (McNeil et al., 2001) scutellar bristle, with lark^{ZF.T:Ivir\HA1} (McNeil et al., 2001) scutellar bristle \| ectopic, with lark^{ZF.T:Ivir\HA1} (McNeil et al., 2001) wing, with lark^{RRM2.RNP2.T:Ivir\HA1} (McNeil et al., 2001) wing, with lark^{RRM2+RTZF.T:Ivir\HA1} (McNeil et al., 2001) wing, with lark^{ZF.T:Ivir\HA1} (McNeil et al., 2001)
Detailed Description
	Statement Reference lark[1] heterozygotes eclose early compared to control flies. No defects in locomotor rhythm are observed when flies expressing lark[dsRNA.Scer\UAS.cSb] under the control of Scer\GAL4[elav-C155] are raised at 25[o]C. However, when the flies are reared at 29[o]C, 21% of flies are arrhythmic and 16% are weakly arrhythmic. Heterozygous lark[1]/+ enhances the phenotype further. No defects in locomotor rhythmicity are observed when flies expressing lark[dsRNA.Scer\UAS.cSb] under the control of Scer\GAL4[nrv2.PS] are raised at 25[o]C. When the flies are reared at 29[o]C, 20% of flies are arrhythmic and 35% are weakly arrhythmic. Heterozygous lark[1]/+ enhances the phenotype further; 42% of these flies are arrhythmic. Only 21% of flies expressing two copies of lark[dsRNA.Scer\UAS.cSb] under the control of Scer\GAL4[tim.Scer\UAS] exhibit robust locomotor rhythms at 29[o]C. This rises to 57% when only one copy of the transgene is expressed. Heterozygous lark[1]/+ enhances the phenotype further; 51% of these flies are completely arrhythmic and a further 29% are weakly arrhythmic. Only 12% of flies expressing two copies of lark[dsRNA.Scer\UAS.cSb] under the control of Scer\GAL4[P2.4.Pdf] (with Dcr-2[Scer\UAS.cDa]) in a heterozygous lark[1] background display robust locomotor rhythms. 37% of flies are arrhythmic. 48% of flies expressing lark[dsRNA.Scer\UAS.cSb] under the control of Scer\GAL4[dimm-929] at 29[o]C exhibit robust locomotor rhythms. The proportion of arrhythmic flies increases when knockdown occurs in a lark[1]/+ mutant background. 40% of flies expressing lark[dsRNA.Scer\UAS.cSb] under the control of Scer\GAL4[Mai179] at 29[o]C exhibit robust locomotor rhythms. The proportion of arrhythmic flies increases when knockdown occurs in a lark[1]/+ mutant background. Only 52% of flies expressing lark[dsRNA.Scer\UAS.cSb] under the control of Scer\GAL4[dimm-929] (restricted to non-pdf expressing cell types using Scer\GAL80[Pdf.PS]) in a lark[1]/+ mutant background at 29[o]C exhibit strong locomotor rhythm defects. Strong locomotor defects are seen in 56% of flies expressing lark[dsRNA.Scer\UAS.cSb] under the control of Scer\GAL4[Mai179] (restricted to non-pdf expressing cell types using Scer\GAL80[Pdf.PS]) in a lark[1]/+ mutant background and raised at 29[o]C. (Sundram et al., 2012) lark¹ animals rescued by lark^{ZF.T:Ivir\HA1} have scutellar bristle defects (such as a missing or malformed posterior scutellar bristle and ectopic bristles). The penetrance of the bristle phenotype is 66-99%. The flies also have a "held out" wing phenotype with 100% penetrance. lark¹ animals rescued by lark^{RRM2.RNP2.T:Ivir\HA1} have scutellar bristle defects (50-100% penetrance) and a "held out" wing phenotype (48-100% penetrance). lark¹ animals rescued by lark^{cMa.T:Ivir\HA1} or lark^{RRM1.RNP2.T:Ivir\HA1} have normal flight behaviour. lark¹ animals rescued by lark^{RRM2+RTZF.T:Ivir\HA1} have scutellar bristle defects (97% penetrance) and an abnormal wing phenotype (94% penetrance). lark¹ homozygotes carrying lark^{cMa.T:Ivir\HA1}, lark^{RRM1.RNP2.T:Ivir\HA1}, lark^{RRM2.RNP2.T:Ivir\HA1} or lark^{ZF.T:Ivir\HA1} show normal eclosion rhythms in 12 hour light:12 hour darkness conditions and normal free-running eclosion rhythms. lark¹ homozygotes carrying lark^{RRM1+RTZF.T:Ivir\HA1} or lark^{RRM2+RTZF.T:Ivir\HA1} have normal free-running eclosion rhythms. (McNeil et al., 2001) Eggs laid by mothers with homozygous lark¹ germline clones have normal morphology, but are extremely fragile, irrespective of whether they are fertilised or not. All embryos arrest during embryonic development, many during syncitial development. Defects in cellularisation and/or tissue differentiation are also seen. (McNeil et al., 1999) P-element has a dominant effect on the timing of eclosion, early eclosion phenotype while entrained to a temperature cycle. Lethality occurs during stage 12 of embryogenesis, the developing nervous system is severely disorganised. (Newby and Jackson, 1993)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhancer of
Statement Reference lark[+]/lark¹ is an enhancer of visible \| dominant phenotype of N^55e11 (Kankel et al., 2007)
Phenotype Manifest In
Enhancer of
Statement Reference lark[+]/lark¹ is an enhancer of wing phenotype of N^55e11 (Kankel et al., 2007)
Additional Comments
Genetic Interactions
Statement Reference Dominantly enhances the Df(1)N-54l9/+ and N[55e11]/+ wing nicking phenotypes. (Kankel et al., 2007)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Rescued by	lark¹ is rescued by lark^+mMa (McNeil et al., 1999) lark¹ is rescued by lark^{cMa.T:Ivir\HA1} (McNeil et al., 1999) lark¹ is rescued by lark^{RRM1.RNP2.T:Ivir\HA1} (McNeil et al., 2001)
Partially rescued by	lark¹ is partially rescued by lark^{RRM1+RTZF.T:Ivir\HA1} (McNeil et al., 2001) lark¹ is partially rescued by lark^{RRM2.RNP2.T:Ivir\HA1} (McNeil et al., 2001) lark¹ is partially rescued by lark^{RRM2+RTZF.T:Ivir\HA1} (McNeil et al., 2001) lark¹ is partially rescued by lark^{ZF.T:Ivir\HA1} (McNeil et al., 2001, McNeil et al., 1999)
Not rescued by	lark¹ is not rescued by lark^{RRM1+2.RNP2.T:Ivir\HA1} (McNeil et al., 2001) lark¹ is not rescued by lark^{ΔC.T:Ivir\HA1} (McNeil et al., 2001)
Comments	The addition of lark^+mMa to homozygous lark¹ germline clones rescues the embryonic fragility and lethality phenotypes. (McNeil et al., 1999)
Stocks ( 0 )

Notes on Origin
Discoverer

Comments
	P-element excision indicates that the single insertion of P{lacW} causes both the behavioural and lethal phenotype. (Newby and Jackson, 1993)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 2 )
Reported As
Symbol Synonym	lark¹ (Sundram et al., 2012, Kankel et al., 2007, Schroeder et al., 2003) unnamed (Newby and Jackson, 1993)
Name Synonym
Secondary FlyBase IDs

References ( 8 )
Research paper	Sundram et al., 2012, J. Biol. Rhythms 27(3): 183--195 Cellular Requirements for LARK in the Drosophila Circadian System. [FBrf0218481] Kankel et al., 2007, Genetics 177(4): 2493--2505 Investigating the genetic circuitry of mastermind in Drosophila, a notch signal effector. [FBrf0202064] Schroeder et al., 2003, J. Neurogenet. 17(2-3): 139--169 Cell-specific expression of the lark RNA-binding protein in Drosophila results in morphological and circadian behavioral phenotypes. [FBrf0167797] McNeil et al., 2001, Genetics 159(1): 229--240 Genetic analysis of functional domains within the Drosophila LARK RNA-binding protein. [FBrf0138413] McNeil et al., 1999, Dev. Genet. 25(4): 387--396 Maternal function of a retroviral-type zinc-finger protein is essential for Drosophila development. [FBrf0111974] McNeil et al., 1998, Neuron 20(2): 297--303 A molecular rhythm mediating circadian clock output in Drosophila. [FBrf0100603] Newby and Jackson, 1996, J. Neurobiol. 31(1): 117--128 Regulation of a specific circadian clock output pathway by lark, a putative RNA-binding protein with repressor activity. [FBrf0089760] Newby and Jackson, 1993, Genetics 135(4): 1077--1090 A new biological rhythm mutant of Drosophila melanogaster that identifies a gene with an essential embryonic function. [FBrf0065510]

Allele Dmel\lark1

Allele Dmel\lark¹