Allele Dmel\sens^E2

General Information
Symbol	Dmel\sens^E2	Species	D. melanogaster
Name		FlyBase ID	FBal0098023
Feature type	allele	Associated gene	Dmel\sens

Allele class	amorphic allele - genetic evidence, 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	amorphic allele - genetic evidence (Chandrasekaran and Beckendorf, 2003) loss of function allele (Nolo et al., 2000, Nolo et al., 2001)
Mutagen	ethyl methanesulfonate (Bloomington Drosophila Stock Center, 1999.7.3)
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 Amino acid replacement: ??@. (Wang et al., 2010) G to A transition at coordinate 3L:13393009 (release 5), resulting in a nonsense mutation. (Wang et al., 2010) Nucleotide substitution: G?A. (Wang et al., 2010) Stop codon in the middle of the coding sequence. (Nolo et al., 2000)
Cytology
Phenotypic Data
Phenotypic Class
	increased cell death (Chandrasekaran and Beckendorf, 2003) lethal \| recessive (Bloomington Drosophila Stock Center, 1999.7.3) visible \| cell autonomous \| recessive \| somatic clone (Nolo et al., 2000) visible \| somatic clone (Pi et al., 2004)
Phenotype Manifest In
	chaeta \| cell autonomous \| somatic clone (Nolo et al., 2000) chaeta \| somatic clone (Pi et al., 2004) dorsal row \| somatic clone (Jafar-Nejad et al., 2006) embryonic/larval oenocyte precursor (Witt et al., 2010) embryonic/larval salivary gland (Chandrasekaran and Beckendorf, 2003) eo neuron \| somatic clone \| supernumerary (Jafar-Nejad et al., 2006) external sensory organ precursor cell IIa \| somatic clone (Jafar-Nejad et al., 2006) external sensory organ precursor cell IIb \| somatic clone \| supernumerary (Jafar-Nejad et al., 2006) mechanosensory chaeta \| somatic clone (Acar et al., 2006) medulla \| somatic clone, with Scer\GAL4^sca-109-68, sens^Scer\UAS.cNa (Morey et al., 2008) microchaeta \| cell autonomous \| somatic clone (Nolo et al., 2000) ommatidium \| somatic clone (with sens^E1) (Frankfort et al., 2001) photoreceptor cell R2 \| somatic clone (Frankfort and Mardon, 2004) photoreceptor cell R5 \| somatic clone (Frankfort and Mardon, 2004) photoreceptor cell R7 \| somatic clone (with sens^E1) (Frankfort et al., 2001) photoreceptor cell R8 \| cell non-autonomous \| somatic clone (with sens^E1) (Frankfort et al., 2001) photoreceptor cell R8 \| somatic clone (Frankfort and Mardon, 2004) photoreceptor cell R8 \| somatic clone (with sens^E1) (Frankfort et al., 2001) postorbital bristle \| somatic clone (Acar et al., 2006) thecogen cell \| somatic clone (Jafar-Nejad et al., 2006) tormogen cell \| cell autonomous \| somatic clone (Nolo et al., 2000) tormogen cell \| somatic clone (Jafar-Nejad et al., 2006) trichogen cell \| somatic clone (Jafar-Nejad et al., 2006) trichome of the posterior wing margin \| somatic clone (Jafar-Nejad et al., 2006) ventral row \| somatic clone (Jafar-Nejad et al., 2006)
Detailed Description
	Statement Reference The number of segments containing oenocytes, the number of oenocytes per segment and the total number of oenocytes per embryo are all significantly decreased in sens[E2] mutant embryos as compared to controls. (Witt et al., 2010) sens[E2] homozygous clones in the eye (in which sens[Scer\UAS.cNa] is expressed under the control of Scer\GAL4[sca-109-68] which rescues R8 cell differentiation, but is not expressed during the late phase of R8 axon targeting) show targeting defects in the medulla; thicker columns, which likely indicate mistargeting of R8 axons to the M6 layer are seen, as well as columns in which the R7 axon terminated in the M6 layer while the R8 axon crosses into a neighbouring column. (Morey et al., 2008) sens^E2 mutant clones lose almost all mechanosensory bristles. The number of post-orbital bristles in sens^E2 mutant clones is less than a third of the wild-type number. (Acar et al., 2006) sens^E2/+ flies do not show a wing bristle phenotype. Induction of sens^E2 clones in the wing leads to a loss of mechanosensory bristles in the anterior wing margin and loss of the noninnervated bristles of the posterior wing margin. Sensory precursors within sens^E2 clones are able to divide, but there is a delay in their division compared with wild-type precursors. The resulting mutant sensory clusters contain multiple neurons and an occasional sheath cell but lack a socket cell in 98% of cases. This indicates that there is a pIIa-to-pIIb transformation in these clones followed by a sheath-to-neuron transformation later in the lineage. (Jafar-Nejad et al., 2006) In sens^E2 homozygous somatic clones in the eye disc, photoreceptor R8 differentiates prematurely as a founder R2/R5 cell which then recruits a reduced number of photoreceptors compared to wild-type. In clones in the resulting adult eyes, ommatidia are disrupted, but there are no undifferentiated regions. (Frankfort and Mardon, 2004) Mutant larvae have small salivary glands, about a half to a third of the size of normal salivary glands. IN addition the salivary glands of stage 16 mutant larvae are smaller than stage 13 embryos, suggesting the loss may be progressive. The mutant salivary glands have less cells than wild-type. The salivary placodes appear to be normally specified and a re similar to wild-type placodes in size and cell numbers. IN addition salivary ducts are normal. Cells in the salivary glands undergo ectopic apoptosis from late stage 12 through to stage 14. (Chandrasekaran and Beckendorf, 2003) When somatic clones of sens^E2/sens^E1 in the eye are made in a Minute background, normally constructed ommatidia containing photoreceptors mutant for sens can only form if the R8 photoreceptor had at least one functional copy of sens. Large patches of mutant ommatidia are readily recovered in adults. These clones are disorganised and of variable size. The number of ommatidia and the spacing between them is not changed from the surrounding wild-type tissue. However all ommatidia have a striking similarity: they do not contain a discernable R8 or R7 photoreceptors. (Frankfort et al., 2001) Heterozygotes do not show a loss of wing margin phenotype. (Nolo et al., 2001) Homozygous clones lack bristles, sockets and microchaetae. The phenotype is cell autonomous. (Nolo et al., 2000)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
NOT suppressed by
Statement Reference sens^E2 has visible \| somatic clone phenotype, non-suppressible by Scer\GAL4^Eq1/phyl^{Scer\UAS.T:Hsap\MYC} (Pi et al., 2004)
Suppressor of
Statement Reference sens^E2/sens[+] is a suppressor \| partially of visible phenotype of mir-9a^J22/mir-9a^E39 (Bejarano et al., 2010) sens^E2 is a suppressor \| somatic clone of visible phenotype of Scer\GAL4^ap-md544, sc^Scer\UAS.cHa (Nolo et al., 2000)
NOT Suppressor of
Statement Reference sens^E2/sens[+] is a non-suppressor of increased cell death phenotype of mir-9a^J22 (Bejarano et al., 2010)
Other
Statement Reference pnr[+]/pnr^VX4, sens^E2 has visible \| dominant phenotype (Asmar et al., 2008) pnr[+]/pnr^VX6, sens^E2 has visible \| dominant phenotype (Asmar et al., 2008)
Phenotype Manifest In
Suppressed by
Statement Reference sens^E1/sens^E2 has photoreceptor cell R8 \| somatic clone \| cell non-autonomous phenotype, suppressible by ro^x63 (Frankfort et al., 2001) sens^E2 has embryonic/larval salivary gland phenotype, suppressible \| partially by Df(3L)H99 (Chandrasekaran and Beckendorf, 2003) sens^E2 has embryonic/larval salivary gland phenotype, suppressible \| partially by Df(3L)XR38 (Chandrasekaran and Beckendorf, 2003) sens^E2 has embryonic/larval salivary gland phenotype, suppressible \| partially by W⁰⁵⁰¹⁴ (Chandrasekaran and Beckendorf, 2003) sens^E2 has embryonic/larval salivary gland phenotype, suppressible by Scer\GAL4^arm.PS/BacA\p35^Scer\UAS.cHa (Chandrasekaran and Beckendorf, 2003)
NOT suppressed by
Statement Reference sens^E2 has adult thorax & microchaeta \| somatic clone phenotype, non-suppressible by Scer\GAL4^tub.PU/da^Scer\UAS.cGa (Jafar-Nejad et al., 2006) sens^E2 has chaeta \| somatic clone phenotype, non-suppressible by Scer\GAL4^Eq1/phyl^{Scer\UAS.T:Hsap\MYC} (Pi et al., 2004) sens^E2 has dorsal row \| somatic clone phenotype, non-suppressible by Scer\GAL4^tub.PU/BacA\p35^Scer\UAS.cHa (Jafar-Nejad et al., 2006) sens^E2 has eo neuron \| supernumerary \| somatic clone phenotype, non-suppressible by Scer\GAL4^tub.PU/da^Scer\UAS.cGa (Jafar-Nejad et al., 2006) sens^E2 has tormogen cell \| somatic clone phenotype, non-suppressible by Scer\GAL4^tub.PU/da^Scer\UAS.cGa (Jafar-Nejad et al., 2006) sens^E2 has trichogen cell \| somatic clone phenotype, non-suppressible by Scer\GAL4^tub.PU/da^Scer\UAS.cGa (Jafar-Nejad et al., 2006) sens^E2 has ventral row \| somatic clone phenotype, non-suppressible by Scer\GAL4^tub.PU/BacA\p35^Scer\UAS.cHa (Jafar-Nejad et al., 2006)
Enhancer of
Statement Reference sens^E2/sens[+] is an enhancer of eo neuron phenotype of sc^10-1 (Jafar-Nejad et al., 2006) sens^E2/sens[+] is an enhancer of medial triple row phenotype of sc^10-1 (Jafar-Nejad et al., 2006) sens^E2/sens[+] is an enhancer of tormogen cell phenotype of sc^10-1 (Jafar-Nejad et al., 2006) sens^E2 is an enhancer of photoreceptor cell \| somatic clone phenotype of rho^7M43, ru¹ (Frankfort and Mardon, 2004)
Suppressor of
Statement Reference sens^E2/sens[+] is a suppressor \| partially of anterior postalar bristle phenotype of mir-9a^J22/mir-9a^E39 (Bejarano et al., 2010) sens^E2/sens[+] is a suppressor \| partially of dorsocentral bristle phenotype of mir-9a^J22/mir-9a^E39 (Bejarano et al., 2010) sens^E2/sens[+] is a suppressor \| partially of wing margin phenotype of mir-9a^J22/mir-9a^E39 (Bejarano et al., 2010) sens^E2 is a suppressor \| somatic clone of adult thorax & macrochaeta phenotype of Scer\GAL4^ap-md544, sc^Scer\UAS.cHa (Nolo et al., 2000) sens^E2 is a suppressor \| somatic clone of adult thorax & microchaeta phenotype of Scer\GAL4^ap-md544, sc^Scer\UAS.cHa (Nolo et al., 2000)
NOT Suppressor of
Statement Reference sens^E2/sens[+] is a non-suppressor of wing pouch phenotype of mir-9a^J22/mir-9a^E39 (Bejarano et al., 2010)
Other
Statement Reference Egfr[+]/Egfr^E1, sens^E2 has eye \| somatic clone phenotype (Frankfort and Mardon, 2004) Egfr[+]/Egfr^E1, sens^E2 has ommatidium \| somatic clone phenotype (Frankfort and Mardon, 2004) Egfr^E1, sens^E2 has eye \| somatic clone phenotype (Frankfort and Mardon, 2004) Egfr^E1, sens^E2 has eye phenotype (Frankfort and Mardon, 2004, Frankfort and Mardon, 2004) Egfr^E1, sens^E2 has ommatidium \| somatic clone phenotype (Frankfort and Mardon, 2004) Egfr^E1, sens^E2 has ommatidium phenotype (Frankfort and Mardon, 2004, Frankfort and Mardon, 2004) pnr[+]/pnr^VX4, sens^E2 has dorsocentral bristle phenotype (Asmar et al., 2008) pnr[+]/pnr^VX6, sens^E2 has dorsocentral bristle phenotype (Asmar et al., 2008)
Additional Comments
Genetic Interactions
Statement Reference sens[E2]/+ results in an approximately 35% reduction in the loss of wing margin caused by mir-9a[J22]/mir-9a[E39]. sens[E2]/+ does not rescue the increased cell death seen in the wing discs of mir-9a[J22]/mir-9a[E39] animals. sens[E2]/+ partially suppresses the formation of ectopic dorsocentral and anterior postalar bristles seen in mir-9a[J22]/mir-9a[E39] adults. (Bejarano et al., 2010) sens[E2]/pnr[VX4] double heterozygotes show loss of dorsocentral bristles. sens[E2]/pnr[VX6] double heterozygotes show loss of dorsocentral bristles. (Asmar et al., 2008) sc^10-1; sens^E2/+ flies show a severe loss of stout bristles at the wing margin in comparison to sc^10-1 single mutants, which show a small loss of these bristles. At the pupal level the sc^10-1; sens^E2/+ double mutants are almost completely lacking the neurons that should have developed from the mechanosensory precursors; this is an enhancement of the reduced number of neurons seen in sc^10-1 single mutant pupae. The double mutants also show a greater loss of socket cells than the single mutants. Thoracic clones that express da^Scer\UAS.cGa, under the control of Scer\GAL4^tub, in a sens^E2 background results in clones that lack microchaetae. Sensory organ precursors in these clones generate extra neurons but no shaft or socket cells, which is similar to sens^E2 single mutant clones. (Jafar-Nejad et al., 2006) In rho^7M43; ru¹; sens^E2 triple mutant clones, no photoreceptors differentiate except for a few photoreceptors near the clonal boundary, presumably rescued non-autonomously by neighboring wild-type cell. No rescue of photoreceptor development is seen when these triple mutant clones are made in a ro^x63 homozygous background. Adult eyes containing sens^E2 homozygous somatic clones induced in an Egfr^E1/+ background are smaller than wild-type. Within the clone there are reduced numbers of ommatidia, as well as gaps of tissue between ommatidia, a phenotype similar to that seen in Egfr^E1 homozygotes. (Frankfort and Mardon, 2004) Expression of phyl^{Scer\UAS.T:Hsap\MYC} under the control of Scer\GAL4^Eq1 fails to rescue external sensory organ formation in homozygous sens^E2 clones in the notum. (Pi et al., 2004) When Df(3L)H99 (A deficiency that uncovers grim, rpr and W) is combined with sens^E2, 93% of salivary glands are normal in size (compared with 0% for sens^E2 mutants). The addition of W⁰⁵⁰¹⁴ or Df(3L)XR38 (which removed rpr) partially suppresses this phenotype as well, though the suppression by W⁰⁵⁰¹⁴ is quantitatively and qualitatively weaker than Df(3L)XR38. (Chandrasekaran and Beckendorf, 2003) The addition of ro^x63 suppresses the photoreceptor cell R8 phenotype seen in sens^E2/sens^E1 somatic clones. (Frankfort et al., 2001) Clones of sens^E2 in the thorax suppress the extra micro- and macrochaetae produced by expression of sc^Scer\UAS.cHa under the control of Scer\GAL4^ap-md544. (Nolo et al., 2000)
Xenogenetic Interactions
Statement Reference Expression of BacA\p35^Scer\UAS.cHa, under the control of Scer\GAL4^tub, in sens^E2 clones fails to rescue the wing margin bristle loss. (Jafar-Nejad et al., 2006)
Complementation & Rescue Data
Complements	sens^SX67 (Nolo et al., 2001)
Fails to complement	sens^E1 (Nolo et al., 2001) sens^E53 (Nolo et al., 2001) sens^E54 (Nolo et al., 2001) sens^E58 (Nolo et al., 2001) sens^E69 (Nolo et al., 2001) sens^I235 (Nolo et al., 2001) sens^Ly-1 (Nolo et al., 2001) sens^M256 (Nolo et al., 2001) sens^SX67R12 (Nolo et al., 2001)
Rescued by	sens^E2 is rescued by sens^g.+t (Acar et al., 2006)
Partially rescued by	sens^E2 is partially rescued by sens^1CCg (Acar et al., 2006)
Not rescued by	sens^E2 is not rescued by sens^1CCg (Acar et al., 2006) sens^E2 is not rescued by sens^2CCg (Acar et al., 2006) sens^E2 is not rescued by sens^3CCg (Acar et al., 2006)
Comments	Expression of sens^g.+t is able to rescue the external sensory organs in adult sens^E2 clones, which lose almost all mechanosensory bristles. Expression of sens^1CCg is able to rescue many of the external sensory organs in adult sens^E2 clones, which lose almost all mechanosensory bristles. Expression of sens^3CCg is unable to rescue the external sensory organs in adult sens^E2 clones, which lose almost all mechanosensory bristles. Expression of sens^g.+t is able to rescue loss of post-orbital bristles in sens^E2 clones. Expression of sens^1CCg is able to partially restore the number of post-orbital bristles in sens^E2 clones to approximately two-thirds of the wild-type. Expression of sens^3CCg is unable to rescue the loss of post-orbital bristles in adult sens^E2 clones. Expression of sens^2CCg is unable to rescue the loss of post-orbital bristles in adult sens^E2 clones. Expression of sens^g.+t rescues the lethality of sens^E2 homozygous embryos. Expression of sens^1CCg does not rescue the lethality of sens^E2 homozygous embryos. However, a significant rescue of PNS development is observed. The rescue is however, not complete, as some of the neurons are lost. In addition, the neurons exhibit differentiation, axon guidance, and fasciculation defects. Expression of sens^3CCg does not rescue the lethality of sens^E2 homozygous embryos. sens^3CCg also does not rescue the PNS defects associated with sens^E2. (Acar et al., 2006)
Stocks ( 4 )
Bloomington	5311 sens^E2 red¹ e¹/TM6B, Tb⁺ 40960 y¹ w^; P{UAS-Mmus\Gfi1}3; mwh¹ sens^E2/TM6B, Tb⁺ 39713 y¹ w^; P{UAS-Mmus\Gfi1B}3/CyO; mwh¹ sens^E2/TM6B, Tb⁺
Kyoto	108169 sens^E2 red¹ e¹/TM6B, Tb¹
Notes on Origin
Discoverer	H. Irick
	Induced on: red¹ e¹ (Bloomington Drosophila Stock Center, 1999.7.3)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 5 )
Reported As
Symbol Synonym	E2 (Nolo et al., 2000) l(3)70Ad^E2 (Bloomington Drosophila Stock Center, 1999.7.3, ) Ly^E2 Ly^sens-E2 sens^E2 (Venken, 2006, Jafar-Nejad et al., 2006, Acar et al., 2006, Christensen et al., 2008.9.29, Zhai et al., 2009, Morey et al., 2008, Bejarano et al., 2010, Li-Kroeger et al., 2008, Witt et al., 2010, Asmar et al., 2008, Wang et al., 2010)
Name Synonym
Secondary FlyBase IDs

References ( 20 )
Research paper	Bejarano et al., 2010, Dev. Biol. 338(1): 63--73 miR-9a prevents apoptosis during wing development by repressing Drosophila LIM-only. [FBrf0209786] Wang et al., 2010, Genome Res. 20(7): 981--988 Rapid identification of heterozygous mutations in Drosophila melanogaster using genomic capture sequencing. [FBrf0211197] Witt et al., 2010, Dev. Biol. 344(2): 1060--1070 Atonal, Senseless, and Abdominal-A regulate rhomboid enhancer activity in abdominal sensory organ precursors. [FBrf0211413] Zhai et al., 2009, Gene Expr. Patterns 9(5): 357--363 Expression of the apoptosis gene reaper in homeotic, segmentation and other mutants in Drosophila. [FBrf0208645] Asmar et al., 2008, Dev. Biol. 316(2): 487--497 Drosophila dLMO-PA isoform acts as an early activator of achaete/scute proneural expression. [FBrf0205286] Li-Kroeger et al., 2008, Dev. Cell 15(2): 298--308 Hox and senseless antagonism functions as a molecular switch to regulate EGF secretion in the Drosophila PNS. [FBrf0205665] Acar et al., 2006, Development 133(10): 1979--1989 Senseless physically interacts with proneural proteins and functions as a transcriptional co-activator. [FBrf0190268] Jafar-Nejad et al., 2006, Development 133(9): 1683--1692 Senseless and Daughterless confer neuronal identity to epithelial cells in the Drosophila wing margin. [FBrf0190299] Frankfort and Mardon, 2004, Development 131(3): 563--570 Senseless represses nuclear transduction of Egfr pathway activation. [FBrf0167533] Pi et al., 2004, Proc. Natl. Acad. Sci. U.S.A. 101(22): 8378--8383 phyllopod is a target gene of proneural proteins in Drosophila external sensory organ development. [FBrf0179385] Chandrasekaran and Beckendorf, 2003, Development 130(19): 4719--4728 senseless is necessary for the survival of embryonic salivary glands in Drosophila. [FBrf0162078] Jafar-Nejad et al., 2003, Genes Dev. 17(23): 2966--2978 Senseless acts as a binary switch during sensory organ precursor selection. [FBrf0167609] Lai, 2003, Genetics 163(4): 1413--1425 Drosophila Tufted is a gain-of-function allele of the proneural Gene amos. [FBrf0158982] Frankfort et al., 2001, Neuron 32(3): 403--414 senseless repression of rough is required for R8 photoreceptor differentiation in the developing Drosophila eye. [FBrf0141686] Nolo et al., 2001, Genetics 157(1): 307--315 Drosophila Lyra mutations are gain-of-function mutations of senseless. [FBrf0132337] Nolo et al., 2000, Cell 102(3): 349--362 Senseless, a Zn finger transcription factor, is necessary and sufficient for sensory organ development in Drosophila. [FBrf0129996]
Supplementary material	Morey et al., 2008, Nature 456(7223): Supplementary Information. [FBrf0209766] Venken, 2006, Science 314(5806): Supporting Online Material. [FBrf0199318]
Personal communication to FlyBase	Christensen et al., 2008.9.29, Isolation and characterization of Df(3L)BSC614. Isolation and characterization of Df(3L)BSC614. [FBrf0205919] Bloomington Drosophila Stock Center, 1999.7.3, Irick and Cherbas Ly region complementation groups. Irick and Cherbas Ly region complementation groups. [FBrf0108618]

Allele Dmel\sensE2

Allele Dmel\sens^E2