Allele Dmel\ems³

General Information
Symbol	Dmel\ems³	Species	D. melanogaster
Name		FlyBase ID	FBal0003714
Feature type	allele	Associated gene	Dmel\ems
Also Known As	ems^9Q64, ems^9Q

Map ( GBrowse )
Allele class	amorphic allele - genetic evidence
Mutagen	ethyl methanesulfonate
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	amorphic allele - genetic evidence (Cohen and Jurgens, 1990)
Mutagen	ethyl methanesulfonate
Mutations Mapped to the Genome

Type Location Additional Notes References point mutation 3R:9,733,321..9,733,321 comment=Site of nucleotide substitution in mutant inferred by FlyBase based on reported amino acid change. evidence=experimental na_change=C9733321T pr_change=Q141@\|ems-P1 reported_pr_change=Q141@ (Dalton et al., 1989)
Associated Sequence Data
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name

UniProtKB/Swiss-Prot
UniProtKB/TrEMBL

Progenitor genotype
Nature of the lesion	Statement Reference stop codon at residue 141
Cytology
Phenotypic Data
Phenotypic Class
	increased cell death \| somatic clone (Lichtneckert et al., 2007) neuroanatomy defective \| adult stage \| somatic clone (Lichtneckert et al., 2008) neuroanatomy defective \| larval stage \| somatic clone (Lichtneckert et al., 2008) neuroanatomy defective \| somatic clone (Das et al., 2008)
Phenotype Manifest In
	adult antennal lobe projection neuron VA1v adPN \| somatic clone (Lichtneckert et al., 2008) adult antennal lobe projection neuron VA3 adPN \| somatic clone (Lichtneckert et al., 2008) adult antennal lobe projection neuron VM2 adPN \| somatic clone (Lichtneckert et al., 2008) adult brain \| somatic clone (Lichtneckert et al., 2007) antennal lobe \| somatic clone (Das et al., 2008) antennal lobe projection neuron adPN \| somatic clone (Lichtneckert et al., 2008) antennal lobe projection neuron lPN \| somatic clone (Lichtneckert et al., 2008) antennal segment (Schmidt-Ott et al., 1994) antennal sense organ (Schmidt-Ott et al., 1994) associated organ (Schmidt-Ott et al., 1995, Schmidt-Ott et al., 1994) Bolwig organ (Schmidt-Ott et al., 1995, Schmidt-Ott et al., 1994) dorsomedial papilla (Schmidt-Ott et al., 1995, Schmidt-Ott et al., 1994) embryonic/larval brain \| somatic clone (Lichtneckert et al., 2007) embryonic/larval optic lobe \| embryonic stage (Schmidt-Ott et al., 1995) fascicle \| somatic clone (Lichtneckert et al., 2007) filzkorper (Hu and Castelli-Gair, 1999) hypocerebral ganglion (Schmidt-Ott et al., 1995) intercalary segment (Schmidt-Ott et al., 1994) local interneuron of larval antennal lobe \| somatic clone (Das et al., 2008) mandibular segment (Schmidt-Ott et al., 1994) neurite \| cell autonomous \| somatic clone (Lichtneckert et al., 2007) neuroblast \| somatic clone (Lichtneckert et al., 2007, Das et al., 2008) ocular segment (Schmidt-Ott et al., 1994) spiracular chamber (Hu and Castelli-Gair, 1999) stigmatophore (Hu and Castelli-Gair, 1999)
Detailed Description
	Statement Reference ems[3] mutant MARCM clones exhibit neuronal loss specifically in the lateral neuroblasts in the 1Nb lineage. This results in a marked reduction in the size of the antennal lobes. (Das et al., 2008) Homozygous and wild-type anterodorsal projection neuron (adPN) clones are recovered with equal frequency (clones induced in early first instar larvae and analysed in late third instar larvae). The average number of cells in homozygous and wild-type adPN clones is virtually identical. Homozygous lateral projection neuron (lPN) clones are recovered at a much lower frequency than wild-type lPN clones (in 2% versus 25% of brains) (clones induced in early first instar larvae and analysed in late third instar larvae). Homozygous lPN clones are much smaller in size (containing fewer cells) than wild-type lPN clones. Homozygous adPN neuroblast clones (induced in the early first instar larva and analysed in the adult brain) show normal cell body projection and axonal projection trajectory, but show marked defects in dendritic targeting, with three types of phenotype being observed. Firstly, mutant adPNs fail to innervate specific glomeruli that are always innervated by wild-type adPNs; the VA1lm glomerulus is innervated by mutant adPNS in only 63% of clones, VA3 in only 71% of clones and VM2 in only 83% of clones. Secondly, mutant adPNs are seen to ectopically innervate inappropriate glomeruli; the DL2 glomerulus is ectopically innervated in 71% of clones, DA2 in 29%, VA6 in 21%, DL5 in 21%, VL2 in 54%, VM1 in 17%, DA1 in 37%, DA4 in 54% and DC1 in 42%. Thirdly, mutant adPNs form inappropriate misprojections into the subesophageal ganglion (this phenotype is seen in approximately one-third of the mutant clones). The axons of homozygous adPN neuroblast clones (induced in the early larva and analysed in the adult) project a fascicle to the lateral horn and form two arborization areas comparable to those of the wild type. Single cell homozygous clones of DL1-innervating adPNs correctly innervate the DL1 glomerulus, project their axons to the lateral horn, bifurcate and form two wild-type-like terminal processes. (Lichtneckert et al., 2008) ems³ mutant brain neuroblast clones induced during the early first instar contain a similar number of cells to wild-type clones at 48 hours after larval hatching (ALH). However, at 72 hours ALH the ems³ mutant clones contain fewer cells than wild-type clones. This difference increase at 96 hours ALH and remains large throughout pupal development and in the adult. BrdU staining shows that the decrease in cell number in ems³ clones is not due to reduced cell proliferation. When examined in the adult brain, many ems³ mutant clones lack the prominent protocerebral fascicle that projects to the superior medial protocerebrum in the wild-type control. In other clones, a reduced protocerebral fascicle is formed. In all clones examined, aberrant projections extend without obvious pattern towards adjacent neuropiles. Misdirected projections of this type are also present in larval ems³ clones. Studies of single cell ems³ MARCM clones shows that these cells fail to extend correct neurite projections and therefore have a cell-autonomous requirement for ems. (Lichtneckert et al., 2007) The stigmatophore protrudes normally but the spiracular chamber is defective in that it lacks a filzkorper and is not connected to the trachea. The stigma in developing embryos fails to slide posteriorly, a phenotype similar to that seen in Abd-B^M1 mutants. The spiracle cells that are in contact with the trachea fail to invaginate, perhaps causing the failure of posterior sliding of the developing stigmatophore. (Hu and Castelli-Gair, 1999) Scr^C1 Antp^Ns-rvC3 Ubx^MX2 abd-A^M1 Abd-B^M8 larvae (deficient for activity of thoracic and abdominal homeotic genes) exhibit sclerotic plates (sp) anterior to each denticle belt. Differentiation of sp depends on ems⁺ function. (Macias and Morata, 1996) The Bolwig organ, dorso-medial and lateral papillae, dorsal organ and associated organ are absent in homozygous embryos. The oesophageal ganglion is reduced in size. The optic lobes are reduced in size and are found in a posterior instead of ventral position in older embryos. (Schmidt-Ott et al., 1995) Mandibular segment, intercalary segments, antennal segments, associated organ, Bolwig's organ, dorsomedial papilla, antennal sense organ and dorsolateral papilla are deleted. Stomatogastric nervous system is present. (Schmidt-Ott et al., 1994) In Ubx^hs.PG embryos that are homozygous for ems³ the C1 belt is very reduced in size. (Gonzalez-Reyes and Morata, 1991) Embryos have no Dll expression in the antennae primordium, it is expressed in other limb primordium. Intercalary and antennal segments and a portion of the pre-antennal region defined by the en head spot are absent in ems mutant embryos. (Cohen and Jurgens, 1990)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Suppressed by
Statement Reference ems³ has increased cell death \| somatic clone phenotype, suppressible by Scer\GAL4^αTub84B.PL/BacA\p35^Scer\UAS.cHa (Lichtneckert et al., 2007) ems³ has neuroanatomy defective \| somatic clone \| adult stage phenotype, suppressible \| partially by acj6^1,4.Scer\UAS/Scer\GAL4^GH146 (Lichtneckert et al., 2008) ems³ has neuroanatomy defective \| somatic clone phenotype, suppressible \| partially by Scer\GAL4^αTub84B.PL/BacA\p35^Scer\UAS.cHa (Lichtneckert et al., 2008)
Phenotype Manifest In
Suppressed by
Statement Reference ems³ has adult antennal lobe projection neuron VA1v adPN \| somatic clone phenotype, suppressible by acj6^1,4.Scer\UAS/Scer\GAL4^GH146 (Lichtneckert et al., 2008) ems³ has antennal lobe projection neuron lPN \| somatic clone phenotype, suppressible \| partially by Scer\GAL4^αTub84B.PL/BacA\p35^Scer\UAS.cHa (Lichtneckert et al., 2008) ems³ has neuroblast \| somatic clone phenotype, suppressible by Scer\GAL4^αTub84B.PL/BacA\p35^Scer\UAS.cHa (Lichtneckert et al., 2007)
NOT suppressed by
Statement Reference ems³ has adult antennal lobe projection neuron VA3 adPN \| somatic clone phenotype, non-suppressible by acj6^1,4.Scer\UAS/Scer\GAL4^GH146 (Lichtneckert et al., 2008) ems³ has adult antennal lobe projection neuron VM2 adPN \| somatic clone phenotype, non-suppressible by acj6^1,4.Scer\UAS/Scer\GAL4^GH146 (Lichtneckert et al., 2008)
Additional Comments
Genetic Interactions
Statement Reference Expression of acj6[1,4.Scer\UAS] under the control of Scer\GAL4[GH146] in homozygous ems[3] adPN neuroblast clones (induced in the early larva and analysed in the adult) significantly rescues the innervation defects of the VA1lm glomerulus, but does not significantly rescue the innervation defects of the VA3 and VM2 glomeruli which are seen in single mutant homozygous ems[3] adPN neuroblast clones. (Lichtneckert et al., 2008)
Xenogenetic Interactions
Statement Reference The reduced recovery of ems[3] lateral projection neuron (lPN) clones is rescued to a wild-type recovery rate if the clones are also expressing BacA\p35[Scer\UAS.cHa] under the control of Scer\GAL4[αTub84B.PL]. The average number of cells in these rescued lPN clones corresponds to 71% of the cell number seen in wild-type clones. (Lichtneckert et al., 2008) Blocking cell death in ems³ neuronal clones by expression of BacA\p35^Scer\UAS.cHa under the control of Scer\GAL4^αTub84B.PL suppresses the low cell number in these clones. (Lichtneckert et al., 2007)
Complementation & Rescue Data
Rescued by	ems³ is rescued by ems^Scer\UAS.cRa/Scer\GAL4^αTub84B.PL (Lichtneckert et al., 2007)
Comments
Stocks ( 0 )

Notes on Origin
Discoverer	Jurgens, Wieschaus, Nusslein-Volhard and Kluding, 1984.

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 4 )
Reported As
Symbol Synonym	ems³ ems^9G (Gonzalez-Reyes and Morata, 1991) ems^9Q64 (Herranz and Morata, 2001, Hu and Castelli-Gair, 1999, Crozatier et al., 1996, Macias and Morata, 1996, Schmidt-Ott et al., 1995, Schmidt-Ott et al., 1994, Dalton et al., 1989, Lichtneckert et al., 2007, Lichtneckert et al., 2008, Hartmann et al., 2010, Das et al., 2008) ems^9Q (Hartmann et al., 2000, Cohen and Jurgens, 1990)
Name Synonym
Secondary FlyBase IDs

References ( 15 )
Research paper	Hartmann et al., 2010, Dev. Biol. 340(1): 125--133 Coral emx-Am can substitute for Drosophila empty spiracles function in head, but not brain development. [FBrf0210189] Das et al., 2008, Neural Dev. 3: 33 Drosophila olfactory local interneurons and projection neurons derive from a common neuroblast lineage specified by the empty spiracles gene. [FBrf0207001] Lichtneckert et al., 2008, Development 135(14): 2415--2424 empty spiracles is required for the development of olfactory projection neuron circuitry in Drosophila. [FBrf0205118] Lichtneckert et al., 2007, Development 134(7): 1291--1300 Cell lineage-specific expression and function of the empty spiracles gene in adult brain development of Drosophila melanogaster. [FBrf0192514] Herranz and Morata, 2001, Development 128(23): 4837--4846 The functions of pannier during Drosophila embryogenesis. [FBrf0141498] Hartmann et al., 2000, Mech. Dev. 90(2): 143--153 Expression, regulation and function of the homeobox gene empty spiracles in brain and ventral nerve cord development of Drosophila. [FBrf0125188] Hu and Castelli-Gair, 1999, Dev. Biol. 214(1): 197--210 Study of the posterior spiracles of Drosophila as a model to understand the genetic and cellular mechanisms controlling morphogenesis. [FBrf0111388] Wiellette et al., 1999, Development 126(23): 5373--5385 spen encodes an RNP motif protein that interacts with Hox pathways to repress the development of head-like sclerites in the Drosophila trunk. [FBrf0112078] Crozatier et al., 1996, Curr. Biol. 6(6): 707--718 collier, a novel regulator of Drosophila head development, is expressed in a single mitotic domain. [FBrf0088068] Macias and Morata, 1996, EMBO J. 15(2): 334--343 Functional hierarchy and phenotypic suppression among Drosophila homeotic genes: The labial and empty spiracles genes. [FBrf0086531] Schmidt-Ott et al., 1995, Rouxs Arch. Dev. Biol. 205(1-2): 31--44 Analysis of neural elements in head-mutant Drosophila embryos suggests segmental origin of the optic lobes. [FBrf0084345] Schmidt-Ott et al., 1994, Proc. Natl. Acad. Sci. U.S.A. 91(18): 8363--8367 Number, identity, and sequence of the Drosophila head segments as revealed by neural elements and their deletion patterns in mutants. [FBrf0075072] Gonzalez-Reyes and Morata, 1991, Development 113(4): 1459--1471 Organization of the Drosophila head as revealed by the ectopic expression of the Ultrabithorax product. [FBrf0053770] Cohen and Jurgens, 1990, Nature 346: 482--485 Mediation of Drosophila head development by gap-like segmentation genes. [FBrf0052647] Dalton et al., 1989, Genes Dev. 3: 1940--1956 Expression and embryonic function of empty spiracles: a Drosophila homeobox gene with two patterning functions on the anterior-posterior axis of the embryo. [FBrf0049810]

Allele Dmel\ems3

Allele Dmel\ems³