Allele Dmel\Dr^Δ68

General Information
Symbol	Dmel\Dr^Δ68	Species	D. melanogaster
Name		FlyBase ID	FBal0063864
Feature type	allele	Associated gene	Dmel\Dr
Also Known As	msh^Δ68

Allele class
Mutagen	P-element activity
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class
Mutagen	P-element activity (Isshiki et al., 1997)
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	P{lacZ-un1}Dr^rH96 (Isshiki et al., 1997)
Nature of the lesion	Statement Reference Imprecise excision of the P{lacZ} element, resulting in a deletion of approximately 4kb that removes a portion of the Dr coding region. (Isshiki et al., 1997)
Cytology
Phenotypic Data
Phenotypic Class
	lethal \| embryonic stage \| recessive (Isshiki et al., 1997) Minute (Villa-Cuesta and Modolell, 2005) mitotic cell cycle defective (Villa-Cuesta and Modolell, 2005) neuroanatomy defective \| embryonic stage (Urbach et al., 2006) neuroanatomy defective \| embryonic stage 12 (Sprecher and Hirth, 2006) neuroanatomy defective \| embryonic stage 15 (Sprecher and Hirth, 2006) partially lethal - majority die (Villa-Cuesta and Modolell, 2005) visible \| recessive \| somatic clone (Milan et al., 2001) visible \| somatic clone (Villa-Cuesta and Modolell, 2005)
Phenotype Manifest In
	abdominal 1 dorsal oblique muscle 1 (Nose et al., 1998) abdominal 1 lateral transverse muscle 1 (Nose et al., 1998) abdominal 1 lateral transverse muscle 2 (Nose et al., 1998) abdominal 1 lateral transverse muscle 3 (Nose et al., 1998) abdominal 1 lateral transverse muscle 4 (Nose et al., 1998) abdominal 1 ventral acute muscle 1 (Nose et al., 1998) abdominal 1 ventral acute muscle 2 (Nose et al., 1998) abdominal 1 ventral acute muscle 3 (Nose et al., 1998) abdominal 2 dorsal oblique muscle 1 (Nose et al., 1998) abdominal 2 dorsal oblique muscle 2 (Nose et al., 1998) abdominal 2 lateral transverse muscle 1 (Nose et al., 1998) abdominal 2 lateral transverse muscle 2 (Nose et al., 1998) abdominal 2 lateral transverse muscle 3 (Nose et al., 1998) abdominal 2 lateral transverse muscle 4 (Nose et al., 1998) abdominal 2 ventral acute muscle 1 (Nose et al., 1998) abdominal 2 ventral acute muscle 2 (Nose et al., 1998) abdominal 2 ventral acute muscle 3 (Nose et al., 1998) abdominal 3 dorsal oblique muscle 1 (Nose et al., 1998) abdominal 3 dorsal oblique muscle 2 (Nose et al., 1998) abdominal 3 lateral transverse muscle 1 (Nose et al., 1998) abdominal 3 lateral transverse muscle 2 (Nose et al., 1998) abdominal 3 lateral transverse muscle 3 (Nose et al., 1998) abdominal 3 lateral transverse muscle 4 (Nose et al., 1998) abdominal 3 ventral acute muscle 1 (Nose et al., 1998) abdominal 3 ventral acute muscle 2 (Nose et al., 1998) abdominal 3 ventral acute muscle 3 (Nose et al., 1998) abdominal 4 dorsal oblique muscle 1 (Nose et al., 1998) abdominal 4 dorsal oblique muscle 2 (Nose et al., 1998) abdominal 4 lateral transverse muscle 1 (Nose et al., 1998) abdominal 4 lateral transverse muscle 2 (Nose et al., 1998) abdominal 4 lateral transverse muscle 3 (Nose et al., 1998) abdominal 4 lateral transverse muscle 4 (Nose et al., 1998) abdominal 4 ventral acute muscle 1 (Nose et al., 1998) abdominal 4 ventral acute muscle 2 (Nose et al., 1998) abdominal 4 ventral acute muscle 3 (Nose et al., 1998) abdominal 5 dorsal oblique muscle 1 (Nose et al., 1998) abdominal 5 dorsal oblique muscle 2 (Nose et al., 1998) abdominal 5 lateral transverse muscle 1 (Nose et al., 1998) abdominal 5 lateral transverse muscle 2 (Nose et al., 1998) abdominal 5 lateral transverse muscle 3 (Nose et al., 1998) abdominal 5 lateral transverse muscle 4 (Nose et al., 1998) abdominal 5 ventral acute muscle 1 (Nose et al., 1998) abdominal 5 ventral acute muscle 2 (Nose et al., 1998) abdominal 5 ventral acute muscle 3 (Nose et al., 1998) abdominal 6 dorsal oblique muscle 1 (Nose et al., 1998) abdominal 6 dorsal oblique muscle 2 (Nose et al., 1998) abdominal 6 lateral transverse muscle 1 (Nose et al., 1998) abdominal 6 lateral transverse muscle 2 (Nose et al., 1998) abdominal 6 lateral transverse muscle 3 (Nose et al., 1998) abdominal 6 lateral transverse muscle 4 (Nose et al., 1998) abdominal 6 ventral acute muscle 1 (Nose et al., 1998) abdominal 6 ventral acute muscle 2 (Nose et al., 1998) abdominal 6 ventral acute muscle 3 (Nose et al., 1998) abdominal 7 dorsal oblique muscle 1 (Nose et al., 1998) abdominal 7 dorsal oblique muscle 2 (Nose et al., 1998) abdominal 7 lateral transverse muscle 1 (Nose et al., 1998) abdominal 7 lateral transverse muscle 2 (Nose et al., 1998) abdominal 7 lateral transverse muscle 3 (Nose et al., 1998) abdominal 7 lateral transverse muscle 4 (Nose et al., 1998) abdominal 7 ventral acute muscle 1 (Nose et al., 1998) abdominal 7 ventral acute muscle 2 (Nose et al., 1998) abdominal 7 ventral acute muscle 3 (Nose et al., 1998) alula \| dorsal \| somatic clone (Milan et al., 2001) basal procephalic longitudinal glia cluster \| embryonic stage 12 (Sprecher and Hirth, 2006) chaeta \| ectopic \| somatic clone (Villa-Cuesta and Modolell, 2005) deutero/tritocerebral embryonic fiber tract founder cluster \| embryonic stage 12 (Sprecher and Hirth, 2006) embryonic/larval cervical connective \| embryonic stage 15 (Sprecher and Hirth, 2006) embryonic/larval subesophageal ganglion \| embryonic stage 15 (Sprecher and Hirth, 2006) embryonic/larval tritocerebral commissure \| embryonic stage 15 (Sprecher and Hirth, 2006) embryonic glial cell & embryonic deutocerebrum & stage 12 embryo (Urbach et al., 2006) embryonic glial cell & embryonic tritocerebrum & stage 12 embryo (Urbach et al., 2006) embryonic tritocerebrum \| embryonic stage 12 (Sprecher and Hirth, 2006) embryonic tritocerebrum \| embryonic stage 15 (Sprecher and Hirth, 2006) frontal connective \| embryonic stage 15 (Sprecher and Hirth, 2006) medial cell body glial cell (Isshiki et al., 1997) medial-most cell body glial cell (Isshiki et al., 1997) mesothoracic dorsal oblique muscle 1 (Nose et al., 1998) mesothoracic dorsal oblique muscle 2 (Nose et al., 1998) mesothoracic lateral transverse muscle 1 (Nose et al., 1998) mesothoracic lateral transverse muscle 2 (Nose et al., 1998) mesothoracic lateral transverse muscle 3 (Nose et al., 1998) mesothoracic lateral transverse muscle 4 (Nose et al., 1998) metathoracic dorsal oblique muscle 1 (Nose et al., 1998) metathoracic dorsal oblique muscle 2 (Nose et al., 1998) metathoracic lateral transverse muscle 1 (Nose et al., 1998) metathoracic lateral transverse muscle 2 (Nose et al., 1998) metathoracic lateral transverse muscle 3 (Nose et al., 1998) metathoracic lateral transverse muscle 4 (Nose et al., 1998) metathoracic ventral acute muscle 1 (Nose et al., 1998) neuroblast NB6-4 (Isshiki et al., 1997) neuron \| embryonic stage (Isshiki et al., 1997) neuropil associated CNS glial cell \| embryonic stage 15 (Sprecher and Hirth, 2006) presumptive embryonic/larval central nervous system \| embryonic stage 15 (Sprecher and Hirth, 2006) prothoracic dorsal oblique muscle 1 (Nose et al., 1998) prothoracic dorsal oblique muscle 2 (Nose et al., 1998) scutellum \| somatic clone (Villa-Cuesta and Modolell, 2005) scutum \| somatic clone (Villa-Cuesta and Modolell, 2005) segment border muscle (Jagla et al., 1999) sensillum campaniformium of proximal dorsal radius \| somatic clone (Villa-Cuesta and Modolell, 2005) tegula \| somatic clone (Villa-Cuesta and Modolell, 2005) wing \| dorsal \| somatic clone (Milan et al., 2001) wing \| somatic clone (Villa-Cuesta and Modolell, 2005) wing hinge \| somatic clone (Villa-Cuesta and Modolell, 2005) wing margin \| anterior \| dorsal \| somatic clone (Milan et al., 2001) wing sclerite \| somatic clone (Villa-Cuesta and Modolell, 2005) wing vein L2 \| dorsal \| somatic clone (Milan et al., 2001) wing vein L3 \| dorsal \| somatic clone (Milan et al., 2001) wing vein L4 \| dorsal \| somatic clone (Milan et al., 2001) wing vein L5 \| dorsal \| somatic clone (Milan et al., 2001)
Detailed Description
	Statement Reference 70% of stage 15 Dr[Δ68] mutant embryos show defects in brain development. These defects range from a gap between the deutocerebral brain region and the neuromeres of the posterior subesophageal ganglion to a reduction in neuronal tissue in the tritocerebrum. Severe axonal pattering disruption is seen in Dr[Δ68] stage 15 embryos. Cervical connectives that run from the deutocerebral and tritocerebral neuromeres to the subesophageal ganglion are reduced or missing and the tritocerebral commissure is disrupted. Defects are also seen in the longitudinal axon tracts in the subesophageal ganglion. Dr[Δ68] stage 15 embryos show reduction or loss of basal procephalic glia associated with both cervical and longitudinal tracts. Neuropile glial loss is seen in the tritocerebral, protocerebral and deutocerebral neuromeres. The frontal connective is severely reduced or missing whereas the supraesophageal commissure appears normal. Surface and cortex glia are unaffected. In Dr[Δ68] stage 12 embryos there is severe loss of basal procephalic longitudinal glia cells. The deuto-/tritocerebral (D/T) fibre tract founder cluster is decreased in size and tract formation in the tritocerebrum is impaired. No significant change in the amount of proliferation in the tritocerebrum is seen in Dr[Δ68] stage 12 embryos compared to wild type. (Sprecher and Hirth, 2006) Mutant stage 12 embryos show a reduction in the number of glial cells in the deutocerebrum and tritocerebrum compared to wild-type embryos, but the number of glial cells in the protocerebrum is unchanged. (Urbach et al., 2006) Dr^Δ68 mitotic recombination clones induced at 0-24hrs after egg laying are generally lethal. Clones induced at 24-48hrs after egg laying are viable. In the wing-disc derivatives, clone-associated defects are displayed in the wing hinge, the posterior scutum and the scutellum. At the hinge, the most frequently observed anomalies are malformations ranging from small defects such as an outheld wing, to the partial loss of proximal hinge structures, or to even the complete loss of most hinge structures, namely sclerites, and the proximal dorsal radius. In the most extreme cases, the wing is fused to the scutellum and scutum, or is displaced posteriorly. Generally, the tegula is not affected. Clone induction between 48-72hrs after egg laying yields similar results. In addition, approximately 19% of flies with visible clones display ectopic tissue carrying macro- and microchaetae, indicating a notum identity. The ectopic notal tissue appears dorsal to the hinge and contiguous to it. At the notum, when Dr^Δ68 minute⁺ clones are induced at 24-48hrs after egg laying, the most frequent anomalies are a reduction of the scutellum and the appearance of depigmented, naked, and corrugated cuticle in the lateral posterior scutum adjacent to the allula and the hinge. Dr^Δ68 mitotic recombination clones frequently develop extra macrochaetae, mostly in the dorsocentral region of the notum and in the scutellum. Occasionally, these clones also induce nearby wild-type cells to differentiate as chaetae. The clones also suppress extant chaetae, the anterior and posterior supraalar bristles being the most affected, and interfere with the correct formation of the scuto-scutellar suture. In clones that comprise the lateral anterior notum, the anterior and posterior notopleural and the presutural macrochaetae are missing in 70, 10 and 15% of cases, respectively. Dr^Δ68 minute⁺ clones in the dorsal hinge exhibit a shortening of the distance between the sc-positive proneural cluster at dorsal radius, in the hinge, and the anterior postalar cluster, in the lateral notum. The distal and proximal sc-positive clusters of the tegula region also appear to be fused. The fold of the disc, that separates the notum and hinge regions is absent when these regions are mutant for Dr^Δ68. (Villa-Cuesta and Modolell, 2005) Homozygous clones have no aberrant phenotype in the ventral surface of the wing, but result in the transformation of dorsal wing structures into a ventral fate. The dorsal anterior wing margin differentiates ventral bristles (a single row of thin bristles interspersed with chemosensory bristles at every fifth position is seen). The dorsal surface of the alula differentiates bristles, reflecting a transformation to the ventral fate. Veins L2 and L4 are corrugated on the dorsal surface and differentiate three rows of strongly pigmented cells, mimicking the ventral pattern. Veins L3 and L5 differentiate ventral characteristics on the dorsal surface, losing pigmentation and consisting of a single row of aligned cells on the dorsal surface. (Milan et al., 2001) The segmental border muscles are enlarged and almost all Kr-positive lateral muscle precursors (LT2 and LT4) are absent in mutant embryos. (Jagla et al., 1999) Embryos exhibit defects in the formation of dorsal muscles 9 and 10, lateral muscles 21-24 and ventral external muscles 26, 27 and 29. (Nose et al., 1998) Neuroblast NB6-4 forms normally in homozygous embryos, although the timing and pattern of its subsequent divisions to produce the glial cells MM-CBG and M-CBG appear abnormal. Medial migration of MM-CBG and M-CBG is often retarded, or fails to occur, and the morphology of these cells is abnormal. The PM and PQ neurons are often located abnormally and have abnormal morphology. (Isshiki et al., 1997)
External Data
Linkouts
Interactions
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Phenotypic Class
Other
Statement Reference Dlw[+]/Dlw¹, Dr^Δ68 has lethal phenotype (Milan et al., 2001) Dlw[+]/Dlw³, Dr^Δ68 has lethal phenotype (Milan et al., 2001) Dlw[+]/Dlw⁴, Dr^Δ68 has lethal phenotype (Milan et al., 2001) Dlw¹, Dr^Δ68 has lethal phenotype (Milan et al., 2001) Dlw³, Dr^Δ68 has lethal phenotype (Milan et al., 2001) Dlw⁴, Dr^Δ68 has lethal phenotype (Milan et al., 2001)
Phenotype Manifest In
Additional Comments
Genetic Interactions
Statement Reference
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Fails to complement	Dr^lacZ-Δ89 (Isshiki et al., 1997) Dr^Δ60 (Isshiki et al., 1997) Dr^Δ138 (Isshiki et al., 1997)
Rescued by	Dr^Δ68 is rescued by Scer\GAL4^how-24B/Scer\GAL4^how-24B/Dr^Scer\UAS.cIa (Nose et al., 1998)
Comments	Scer\GAL4^how-24B-mediated expression of Dr^Scer\UAS.cIa rescues the lateral muscle phenotype. (Nose et al., 1998)
Stocks ( 0 )

Notes on Origin
Discoverer

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 4 )
Reported As
Symbol Synonym	Dr^Δ68 msh^d68 (Urbach et al., 2006) msh^delta68 (Yagi et al., 1998) msh^Δ68 (Villa-Cuesta and Modolell, 2005, Ashraf and Ip, 2001, Nose et al., 1998, Isshiki et al., 1997, Jagla et al., 1999, Von Ohlen et al., 2009, Müller et al., 2010, Sprecher and Hirth, 2006)
Name Synonym
Secondary FlyBase IDs

References ( 12 )
Research paper	Müller et al., 2010, PLoS ONE 5(12): e14323 Regulation and Functions of the lms Homeobox Gene during Development of Embryonic Lateral Transverse Muscles and Direct Flight Muscles in Drosophila. [FBrf0212632] Von Ohlen et al., 2009, Dev. Dyn. 238(11): 2735--2744 Ind represses msh expression in the intermediate column of the Drosophila neuroectoderm, through direct interaction with upstream regulatory DNA. [FBrf0208969] Sprecher and Hirth, 2006, Dev. Dyn. 235(11): 2920--2929 Expression and function of the columnar patterning gene msh in late embryonic brain development of Drosophila. [FBrf0194471] Urbach et al., 2006, Development 133(21): 4315--4330 Segment-specific requirements for dorsoventral patterning genes during early brain development in Drosophila. [FBrf0194392] Villa-Cuesta and Modolell, 2005, Development 132(18): 4087--4096 Mutual repression between msh and Iro-C is an essential component of the boundary between body wall and wing in Drosophila. [FBrf0187488] Jagla et al., 2002, Development 129(4): 1037--1047 Cross-repressive interactions of identity genes are essential for proper specification of cardiac and muscular fates in Drosophila. [FBrf0144816] Ashraf and Ip, 2001, Development 128(23): 4757--4767 The Snail protein family regulates neuroblast expression of inscuteable and string, genes involved in asymmetry and cell division in Drosophila. [FBrf0141494] Milan et al., 2001, Development 128(17): 3263--3268 msh specifies dorsal cell fate in the Drosophila wing. [FBrf0138371] Jagla et al., 1999, Int. J. Dev. Biol. 43(6): 571--573 Plasticity within the lateral somatic mesoderm of Drosophila embryos. [FBrf0123062] Nose et al., 1998, Development 125(2): 215--223 Regional specification of muscle progenitors in Drosophila: the role of the msh homeobox gene. [FBrf0100612] Yagi et al., 1998, Development 125(18): 3625--3633 Interaction between Drosophila EGF receptor and vnd determines three dorsoventral domains of the neuroectoderm. [FBrf0104795] Isshiki et al., 1997, Development 124(16): 3099--3109 The role of the msh homeobox gene during Drosophila neurogenesis: implication for the dorsoventral specification of the neuroectoderm. [FBrf0098274]

Allele Dmel\DrΔ68

Allele Dmel\Dr^Δ68