Allele Dmel\smo³

General Information
Symbol	Dmel\smo³	Species	D. melanogaster
Name		FlyBase ID	FBal0015766
Feature type	allele	Associated gene	Dmel\smo
Also Known As	smo^Q14, smo3

Map ( GBrowse )
Allele class	cold sensitive amorphic allele - genetic evidence, cold sensitive 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	cold sensitive amorphic allele - genetic evidence (Chen and Struhl, 1996) cold sensitive loss of function allele (Alcedo et al., 1996)
Mutagen	ethyl methanesulfonate (Tearle and Nusslein-Volhard, 1987)
Mutations Mapped to the Genome

Type Location Additional Notes References point mutation 2L:279,368..279,368 comment=TGG to TGA evidence=experimental na_change=G279368A pr_change=W366@\|smo-PA reported_na_change=G1783A reported_pr_change=W366@ (Chen and Struhl, 1998, Alcedo et al., 2000)
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: W366@. Nucleotide substitution: G1783A. (Alcedo et al., 2000) Amino acid replacement: W366@. W366 falls just after the coding sequence for the third transmembrane domain. (Chen and Struhl, 1998)
Cytology
Phenotypic Data
Phenotypic Class
	increased cell death \| somatic clone (Vrailas and Moses, 2006) lethal (with smo^1K) (Nakano et al., 2004) lethal (with smo^2A) (Nakano et al., 2004) lethal (with smo^3J) (Nakano et al., 2004) lethal (with smo^4C2) (Nakano et al., 2004) lethal (with smo^4DI) (Nakano et al., 2004) lethal (with smo^4F4) (Nakano et al., 2004) lethal (with smo^IA3) (Nakano et al., 2004) lethal \| recessive \| segment polarity expression pattern \| cold sensitive (Alcedo et al., 1996) mitotic cell cycle defective \| cell autonomous \| somatic clone (Firth and Baker, 2005) mitotic cell cycle defective \| cell non-autonomous \| somatic clone (Firth and Baker, 2005) mitotic cell cycle defective \| somatic clone (Vrailas and Moses, 2006) visible (Lawrence et al., 1999, Lawrence et al., 1999) visible, with Scer\GAL4^C-765, smo^{5A.Scer\UAS.T:Zzzz\FLAG} (Collins and Cohen, 2005) visible, with Scer\GAL4^ptc-559.1, smo^{dsRNA.Scer\UAS.WIZ} (Casso et al., 2008) visible \| recessive \| somatic clone (Kopp and Duncan, 2002, Amanai and Jiang, 2001, Vegh and Basler, 2003) visible \| somatic clone (Gorfinkiel et al., 1999, Chen and Struhl, 1996, Moreno and Morata, 1999, Sweeney et al., 2007, Christiansen et al., 2012)
Phenotype Manifest In
	1st posterior cell, with Scer\GAL4^C-765, smo^{5A.Scer\UAS.T:Zzzz\FLAG} (Collins and Cohen, 2005) abdominal tergite 8 \| somatic clone (Gorfinkiel et al., 1999) adult abdomen & cuticle \| anterior compartment (Struhl et al., 1997) adult abdomen & microchaeta \| anterior compartment (Struhl et al., 1997) adult cuticle \| somatic clone (Moreno and Morata, 1999) adult head \| somatic clone (Vegh and Basler, 2003) adult thorax \| somatic clone (Vegh and Basler, 2003) anal plate \| somatic clone (Gorfinkiel et al., 1999) clasper \| somatic clone (Gorfinkiel et al., 1999) denticle belt (Gritzan et al., 1999, Quirk et al., 1997) double row \| ectopic \| somatic clone (Vegh and Basler, 2003) embryonic/first instar larval cuticle \| cold sensitive (Alcedo et al., 1996) embryonic epidermis (Ingham et al., 2000, Gritzan et al., 1999) embryonic epidermis \| segment polarity expression pattern \| cold sensitive (Alcedo et al., 1996) epiproct \| somatic clone (Gorfinkiel et al., 1999) eye \| somatic clone (Amanai and Jiang, 2001, Christiansen et al., 2012) eye disc \| cell autonomous \| somatic clone (Firth and Baker, 2005) eye disc \| cell non-autonomous \| somatic clone (Curtiss and Mlodzik, 2000, Firth and Baker, 2005) eye disc \| somatic clone (Greenwood and Struhl, 1999, Vrailas and Moses, 2006) female gonopod \| somatic clone (Gorfinkiel et al., 1999) follicle stem cell \| somatic clone (Kirilly et al., 2005) genital arch \| somatic clone (Gorfinkiel et al., 1999) hypandrial bristle \| somatic clone (Gorfinkiel et al., 1999) lamina & neuron \| precursor \| somatic clone (Huang and Kunes, 1998) leg \| somatic clone (Vegh and Basler, 2003) maxillary palp olfactory receptor neuron \| cell non-autonomous \| somatic clone (Sweeney et al., 2007) maxillary palpus \| somatic clone (Sweeney et al., 2007) medial ocellus \| somatic clone \| supernumerary (Thomas and Ingham, 2003) microchaeta \| ectopic \| somatic clone (Aldaz et al., 2003) morphogenetic furrow (Strutt and Mlodzik, 1997) morphogenetic furrow & nucleus \| somatic clone (D'Costa et al., 2006) ocellus \| somatic clone (Thomas and Ingham, 2003) ommatidial precursor cluster (Dominguez, 1999) photoreceptor cell (Strutt and Mlodzik, 1997) photoreceptor cell & neuron (Dominguez, 1999) photoreceptor cell \| cell non-autonomous \| somatic clone (Curtiss and Mlodzik, 2000) photoreceptor cell R8 & eye disc \| somatic clone (Fu and Baker, 2003) photoreceptor cell R8 \| precursor (Dominguez, 1999) scutellum \| somatic clone (Aldaz et al., 2003) tergite \| anterior \| somatic clone (Kopp and Duncan, 2002) tergite \| anterior compartment (Lawrence et al., 1999, Lawrence et al., 1999) third segment of antenna \| somatic clone (Sweeney et al., 2007) wing (Rodriguez and Basler, 1997) wing \| posterior compartment \| somatic clone (Chen and Struhl, 1996) wing \| somatic clone (Vegh and Basler, 2003) wing disc \| anterior/posterior compartment boundary (Rodriguez and Basler, 1997) wing margin \| somatic clone (Vegh and Basler, 2003) wing pouch \| somatic clone (Schilling et al., 2011) wing vein \| ectopic \| somatic clone (Vegh and Basler, 2003) wing vein L3, with Scer\GAL4^C-765, smo^{5A.Scer\UAS.T:Zzzz\FLAG} (Collins and Cohen, 2005) wing vein L3, with Scer\GAL4^ptc-559.1, smo^{dsRNA.Scer\UAS.WIZ} (Casso et al., 2008) wing vein L4, with Scer\GAL4^C-765, smo^{5A.Scer\UAS.T:Zzzz\FLAG} (Collins and Cohen, 2005) wing vein L4, with Scer\GAL4^ptc-559.1, smo^{dsRNA.Scer\UAS.WIZ} (Casso et al., 2008) wing vein L4 \| ectopic \| somatic clone (Vegh and Basler, 2003) wing vein L4 \| somatic clone (Vegh and Basler, 2003)
Detailed Description
	Statement Reference Homozygous clones in the eye induced using the ey-FLP/FRT system result in eyes that are smaller than normal with over-represented smo[+] tissue. (Christiansen et al., 2012) Somatic clones of homozygous smo[3] mutant cells situated close to the anterior side of the anterior-posterior compartment boundary, but not on the posterior side, can be observed to round off. In addition, a tendency of the smo[3] mutant clones that immediately straddle the boundary to migrate from anterior towards posterior territory (but not in the opposite direction) is observed. (Schilling et al., 2011) The mutant wing vein phenotype produced by expression of smo[dsRNA.Scer\UAS.WIZ] under the control of Scer\GAL4[ptc-559.1] is enhanced by smo[3]; the extent of fusion of veins 3 and 4 is increased. (Casso et al., 2008) Cells in homozygous clones straddling the morphogenetic furrow in the eye disc can constrict apically and shorten along their apicobasal axis and can be part of a morphogenetic furrow, although the furrow in the mutant clone is often more posterior in comparison to neighbouring wild-type cells. (Schlichting and Dahmann, 2008) Homozygous clones generated using the eyFLP technique result in the occasional loss of one or both third antennal segments or maxillary palps. In animals containing homozygous smo[3] clones induced by eyFLP and with two intact antennae, homozygous Or42a-expressing olfactory receptor neurons (ORNs) from the maxillary palp correctly target to their normal glomerulus. However, when both antenna fail to form in animals containing homozygous smo[3] clones, Or42a-expressing maxillary palp ORNs mistarget (7 out of 9 cases), The axons often mistarget to areas normally occupied by antennal ORN axons. Or23a-expressing ORN axons from the antenna project to their normal glomeruli in animals containing homozygous smo[3] clones induced by eyFLP and with no maxillary palp. (Sweeney et al., 2007) In smo³ somatic clones that span the morphogenetic furrow, nuclei remain apical, rather than moving to the basal side of the cell in the furrow as wild-type cells do. (D'Costa et al., 2006) At least some of the cells in smo[3] homozygous somatic clones in the eye disc go on to express markers consistent with normal differentiation of photoreceptors. However, they fail to re-enter the cell cycle at the second mitotic wave. These clones also show evidence of increased cell-death, including dysmorphic nuclei and expression of cell-death markers. (Vrailas and Moses, 2006) smo[3] homozygous clones in the dorsal air sac primordium grow normally and populate the tip of the air sac to the same degree as wild-type clones. (Cabernard and Affolter, 2005) The reduction in distance between wing veins III and IV and partial fusions of these veins proximal to the anterior crossvein that is seen in flies expressing smo^{5A.Scer\UAS.T:Zzzz\FLAG} under the control of Scer\GAL4^C-765 are strongly enhanced by smo³/+. (Collins and Cohen, 2005) G1 arrest is often delayed by 1-2 cell diameters in smo[3] eye disc cells. S phases and mitoses of the second mitotic wave are delayed but not absent in smo[3] mutant cells. However, if the cells immediately posterior are wild type, then smo[3] mutant cells enter S phase at the normal time. (Firth and Baker, 2005) Unlike neutral somatic clones, smo³ homozygous somatic clones are rapidly lost from the somatic stem cell population of the germarium. (Kirilly et al., 2005) smo³ somatic clones in the scutellum differentiate microchaetes, indicative of transformation into scutum. (Aldaz et al., 2003) R8 photoreceptor differentiation is delayed in smo³ mutant somatic clones. (Fu and Baker, 2003) Somatic clones of smo³ in the ocellar triangle (ocellar cuticle) lead to reduced or absent ocelli. The medial ocellus is occasionally split in two. (Thomas and Ingham, 2003) Homozygous clones in the wing induced using Scer\FLP1^hs.PS can result in the disruption or partial duplication of wing vein L4, and it may be shifted posteriorly. In addition, double-row wing margin bristles occasionally appear more posteriorly along the wing margin than normal. More severe phenotypes show an increasing anteriorisation of the wing. Homozygous clones in the wing induced using Scer\FLP1^Scer\UAS.cCa expressed under the control of Scer\GAL4^dpp.blk1, Scer\GAL4^spalt or Scer\GAL4^ptc-559.1 result in ectopic veins appearing between veins L3 and L4, but no displacement of L4 or defects of the bristle pattern are seen. Homozygous clones induced by using Scer\FLP1^Scer\UAS.cCa expressed under the control of Scer\GAL4^dpp.blk1 or Scer\GAL4^ptc-559.1 result in defects in the thorax, head and legs that weaken the flies considerably. Homozygous clones induced by using Scer\FLP1^Scer\UAS.cCa expressed under the control of Scer\GAL4^vg.int2.1 result in a wide variety of adult wing defects, but despite the disruption in wing patter, these flies are fully viable and do not show defects in other tissues. (Vegh and Basler, 2003) Ommatidial rotation is normal in smo³ mutant clones (n = 53). (Chou and Chien, 2002) Homozygous mutant clones of glial cells do not migrate prematurely at second instar and are found exclusively posterior to the morphogenetic furrow. (Hummel et al., 2002) Large homozygous clones in the anterior tergite that abut the segment boundary and span the a1, a2 and a3 regions have completely normal polarity both within the clone and in surrounding wild-type cells. The a1 region shows transformation of the cuticle to a2 cuticle. Small homozygous clones in the anterior tergite entirely within the a1 region, and separated from the a2 region by a strip of untransformed a1 cuticle occasionally (2/13 cases) show altered polarity in 1 or 2 cells along the posterior edge of the clone. (Kopp and Duncan, 2002) Mosaic flies with eyes homozygous for smo³ (clones generated using the "eyFLP" system) have small eyes. (Amanai and Jiang, 2001) When somatic clones of smo³ are made in the developing eye, there is no significant reduction in the percentage of homozygous mutant cells, and the relative spatial distribution of the glia in the eye disc remains unaffected. (Rangarajan et al., 2001) Photoreceptor development is blocked in the central regions of marginal homozygous clones in the eye. This effect is nonautonomous; photoreceptors develop along the outer edges of marginal clones and eventually, throughout internal clones in the eye. (Curtiss and Mlodzik, 2000) Homozygotes embryos exhibit a lawn of denticles covering most of the ventral surface. (Ingham et al., 2000) When analysed in clones in the developing eye the number of R8 cell precursors is reduced and their spacing is irregular. The timing of emergence of the mutant R8 cells is normal. Neuronal cells in clones autonomously suffer delay in differentiation. Packing in the ommatidial clusters is disrupted. (Dominguez, 1999) Early smo³ somatic clones (24-48hr after egg-laying (AEL)) in the female analia delete part of the dorsal anal plate but do not affect the ventral plate. Clones in the abdominal tergite 8 induce overgrowth. Clones in the perianal ring produce duplicated anal structures. The long bristle of the vaginal plates can also be duplicated. Small clones outside the A/P compartment border had no effect. Clones in male adult terminalia autonomously duplicate the structures close to the Antero-posterior border such as the genital arc, the claspers, and the hypandrium bristle. Large clones in the male analia delete most of the anal plate. (Gorfinkiel et al., 1999) Cells in homozygous clones in the eye disc are capable of differentiating as photoreceptors. However, there is a significant delay - there is a clear posterior displacement in the onset of photoreceptor differentiation in mutant cells; differentiation is first seen at the posterior, and occasionally lateral, edges of the clone. (Greenwood and Struhl, 1999) Denticle belts are made up of mostly type 5 denticles. (Gritzan et al., 1999) Clones induced in the embryo in the posterior domain of the A compartment make a3 cuticle. The most posterior clones move back, behind the twin clone. The mutant clone assumes a smooth elliptical shape. In the a3 territory the boundaries of the clone are wiggly. (Lawrence et al., 1999) smo³ clones induced before 72 hours development frequently produce regions of non-cuticular material, thought to be hindgut, in areas that are normally cuticular. (Moreno and Morata, 1999) Homozygous clones that include the posterior margin of the lamina furrow lack S phase lamina neuron precursors (LPCs). The scattered S phase cells anterior to the lamina furrow, ad the distribution of S phase cells in other proliferation centres, such as the outer proliferation centre, are unaffected in homozygous clones. (Huang and Kunes, 1998) Shows a weak dominant enhancing effect on B mutations. (Epps et al., 1997) Homozygous phenotype at 25^oC is highly variable, ranging from almost wild type to the occasional partial fusion of denticle belts. At 18^oC the denticle belts are completely eliminated and the ventral surface of the cuticle is covered in a continuous lawn of denticles of similar size and random polarity. (Quirk et al., 1997) Clones of cells mutant for smo redirect the A/P affinity boundary in the developing wing disc. They form a straight boundary when juxtaposed with sister smo⁺ or smo⁺/smo^- A cells, but a wiggly boundary with neighboring smo^-/smo⁺ cells in the P compartment. Similar results are seen in the adult wing. smo^- cells autonomously form anterior wing margin structures if they are derived from A cells, even when they are located in the domain normally occupied by P-compartment cells. (Rodriguez and Basler, 1997) Clones in the anterior compartment of the adult abdomen develop normally when they arise in the middle of the anterior compartment. Clones that arise near or at the anterior/posterior compartment boundary are transformed, forming cuticle characteristic of a more anterior position within the anterior compartment. This transformation is autonomous. Hairs both within and surrounding the clone often have abnormal polarity, although mutant cells which lie very close to the boundary have hairs with normal polarity. Clones that arise in the a1 anterior region of the anterior compartment are transformed to make a2 cuticle. (Struhl et al., 1997) Clones in the developing eye retard the progression of the morphogenetic furrow. Photoreceptor differentiation is retarded, but not prevented, concomitantly with furrow progression. Clones in the eye that lack both Pka-C1 and smo behave like loss of function Pka-C1 clones. Clones show ectopic photoreceptor differentiation and eventually merge with the endogenous field of differentiation, show no retardation of the furrow, pass through a furrow fate and induce non-autonomous ectopic photoreceptor differentiation outside the clone. (Strutt and Mlodzik, 1997) Mutant embryos show a cold sensitive segment polarity phenotype. At 25^oC segmental defects are mild whereas at 18^oC embryos variably show a classic segment polarity cuticle phenotype. (Alcedo et al., 1996) Homozygous clones in the developing wing that arise distant from the A/P boundary develop normally. Clones that arise immediately anterior to the A/P boundary are associated with reorganised wing patterns to form a mirror symmetric double-anterior winglet and conversely the posterior compartment is severely reduced. (Chen and Struhl, 1996) Does not interact with RpII140^wimp maternal effect. (Parkhurst and Ish-Horowicz, 1991) cold-sensitive
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Suppressor of
Statement Reference smo[+]/smo³ is a suppressor of hyperplasia phenotype of boi^e01708 (Hartman et al., 2010)
Other
Statement Reference Df(2R)en^E, smo³ has visible phenotype (Lawrence et al., 1999, Lawrence et al., 1999) Mad¹², smo³ has decreased cell number \| somatic clone phenotype (Rangarajan et al., 2001, Rangarajan et al., 2001) smo³, tkv⁸ has mitotic cell cycle defective \| somatic clone phenotype (Baonza and Freeman, 2005)
Phenotype Manifest In
Enhanced by
Statement Reference Scer\GAL4^Bx-MS1096, smo³, smo^{C.Scer\UAS.T:Hsap\MYC,T:Uuuu\Myr4} has costal cell phenotype, enhanceable by fu^Scer\UAS.cAb/Scer\GAL4^Bx-MS1096 (Hooper, 2003) Scer\GAL4^Bx-MS1096, smo³, smo^{C.Scer\UAS.T:Hsap\MYC,T:Uuuu\Myr4} has wing vein \| ectopic phenotype, enhanceable by fu^Scer\UAS.cAb/Scer\GAL4^Bx-MS1096 (Hooper, 2003) smo³ has photoreceptor cell R8 & eye disc \| somatic clone phenotype, enhanceable \| somatic clone by Mad^1-2 (Fu and Baker, 2003) smo³ has photoreceptor cell R8 & eye disc \| somatic clone phenotype, enhanceable \| somatic clone by tkv⁸ (Fu and Baker, 2003)
Suppressed by
Statement Reference Pka-C1^E95, smo³ has wing disc \| ectopic phenotype, suppressible by sgg^M11 (Price and Kalderon, 2002) Scer\GAL4^Bx-MS1096, fz::smo^SSF.Scer\UAS, smo[+]/smo³ has 1st posterior cell phenotype, suppressible by Scer\GAL4^Bx-MS1096, fz::smo^SSF.Scer\UAS (Hooper, 2003) Scer\GAL4^Bx-MS1096, smo³, smo^{C.Scer\UAS.T:Hsap\MYC,T:Uuuu\Myr4} has 1st posterior cell phenotype, suppressible \| partially by fu^Scer\UAS.cAb/Scer\GAL4^Bx-MS1096 (Hooper, 2003) smo³ has denticle belt phenotype, suppressible by wg^Scer\UAS.cLa/Scer\GAL4^h-1J3 (Quirk et al., 1997) smo³ has photoreceptor cell R8 & eye disc \| somatic clone phenotype, suppressible \| somatic clone by ci⁹⁴ (Fu and Baker, 2003)
NOT suppressed by
Statement Reference Pka-C1^E95, smo³ has mesothoracic tergum \| ectopic phenotype, non-suppressible by sgg^M11 (Price and Kalderon, 2002) smo³ has follicle stem cell \| somatic clone phenotype, non-suppressible by Scer\GAL4^Act5C.PI/Scer\GAL4^Act5C.PI/tkv^{Q199D.Scer\UAS} (Kirilly et al., 2005) smo³ has phenotype, non-suppressible by eRF1[+]/eRF1^F2 (Chao et al., 2003)
Enhancer of
Statement Reference smo[+]/smo³ is an enhancer of wing phenotype of Moe^C858 (Molnar and de Celis, 2006)
Suppressor of
Statement Reference smo[+]/smo³ is a suppressor of follicle stem cell phenotype of boi^e01708 (Hartman et al., 2010) smo³ is a suppressor of denticle belt phenotype of ptc^G12 (Quirk et al., 1997)
NOT Suppressor of
Statement Reference smo³ is a non-suppressor \| somatic clone of phenotype of en^Apa (Kopp and Duncan, 2002)
Other
Statement Reference Df(2R)en^E, smo³ has pleurum \| posterior compartment phenotype (Lawrence et al., 1999) Df(2R)en^E, smo³ has tergite \| anterior compartment phenotype (Lawrence et al., 1999) Df(2R)en^E, smo³ has tergite \| posterior compartment phenotype (Lawrence et al., 1999) Df(2R)en^E, smo³ has wing disc \| posterior \| somatic clone phenotype (Dahmann and Basler, 2000) ed^1X5, smo³ has wing disc \| somatic clone \| third instar larval stage phenotype (Wei et al., 2005) en^E, inv^E, smo³ has wing disc \| posterior \| somatic clone phenotype (Dahmann and Basler, 2000, Dahmann and Basler, 2000) hh^αTub84B.PB, smo³ has wing disc \| somatic clone phenotype (Dahmann and Basler, 2000, Dahmann and Basler, 2000) Mad^1-2, smo³ has eye disc \| somatic clone phenotype (Curtiss and Mlodzik, 2000, Curtiss and Mlodzik, 2000) Mad^1-2, smo³ has photoreceptor cell \| somatic clone phenotype (Curtiss and Mlodzik, 2000, Curtiss and Mlodzik, 2000) Mad¹², smo³ has eye disc \| somatic clone phenotype (Rangarajan et al., 2001, Rangarajan et al., 2001) Pka-C1^E95, smo³ has mesothoracic tergum \| ectopic phenotype (Price and Kalderon, 2002, Price and Kalderon, 2002) Pka-C1^E95, smo³ has wing disc \| ectopic phenotype (Price and Kalderon, 2002, Price and Kalderon, 2002) Scer\GAL4^Bx-MS1096, fz::smo^SSF.Scer\UAS, smo[+]/smo³ has 1st posterior cell phenotype (Hooper, 2003) Scer\GAL4^Bx-MS1096, fz::smo^SSF.Scer\UAS, smo³ has 1st posterior cell phenotype (Hooper, 2003) slmb¹, smo³ has wing phenotype (Jiang and Struhl, 1998) slmb², smo³ has wing phenotype (Jiang and Struhl, 1998) smo³, tkv⁴ has morphogenetic furrow \| somatic clone phenotype (Schlichting and Dahmann, 2008) smo³, tkv⁸ has eye disc \| somatic clone phenotype (Baonza and Freeman, 2005) smo³, tkv⁸ has photoreceptor cell phenotype (Greenwood and Struhl, 1999, Greenwood and Struhl, 1999)
Additional Comments
Genetic Interactions
Statement Reference The follicle stem cell hyper-proliferation and excess follicle cell accumulation phenotypes are suppressed in boi[e01708] mutants by smo[3]/+. (Hartman et al., 2010) Cells in smo[3] tkv[4] double homozygous clones straddling the morphogenetic furrow fail to constrict apically and do not shorten along their apicobasal axis, failing to form a morphogenetic furrow. Expression of Cad86C[Scer\UAS.T:Ivir\HA1] under the control of Scer\GAL4[Act5C.PP] in smo[3] tkv[4] double homozygous clones in the region of the morphogenetic furrow can result in the formation of an epithelial invagination in some cases. Cells at the centre of this invagination are shorter along their apicobasal axis compared to smo[3] tkv[4] double homozygous clones or to wild-type cells within the morphogenetic furrow. (Schlichting and Dahmann, 2008) The more severe wing defects seen in Moe^c858 homozygotes (veination defects and the formation of vesicles between the dorsal and ventral wing surfaces) are enhanced by smo³/+. (Molnar and de Celis, 2006) Marker analysis shows that some photoreceptor differentiation begins in some fraction of the cells in smo[3] tkv[8] double homozygous cones in the late third instar eye disc. (Vrailas and Moses, 2006) In smo[3], tkv[8] double mutant eye disc clones, BrdU incorporation, a marker of S phase, is abolished in the second mitotic wave. (Baonza and Freeman, 2005) The rapid loss of smo³ homozygous somatic clone cells from the somatic stem cell population of the germarium is not significantly suppressed if the clone cells are also tkv^{Q199D.Scer\UAS}; Scer\GAL4^Act5C.PI (Scer\GAL80 method). (Kirilly et al., 2005) Independent ed^1X5, smo³ double mutant clones that originate in different compartments (as evidenced by ci or en expression) can fuse together to form composite clones which have roundish, smooth shapes. Within such composite clones, the characteristic segregation of 'anterior' and 'posterior' cells is maintained - they do not intermix. (Wei et al., 2005) R8 differentiation fails in tkv⁸; smo³ or Mad^1-2; smo³ double mutant clones, whereas it is only delayed in smo³ single mutant clones. The delay in R8 photoreceptor differentiation seen in smo³ mutant somatic clones is suppressed by ci⁹⁴. (Fu and Baker, 2003) Expression of high levels of fz::smo^SSF.Scer\UAS under the control of Scer\GAL4^Bx-MS1096 in a smo³/+ background results in a reduction in the spacing between the 3rd and 4th wing vein. Co-expression of fu^Scer\UAS.cAb only weakly suppresses the reduction in spacing between the 3rd and 4th wing vein seen in flies expressing high levels of fz::smo^SSF.Scer\UAS under the control of Scer\GAL4^Bx-MS1096 in a smo³/+ background. The reduction in spacing between the 3rd and 4th wing vein seen in flies expressing high levels of fz::smo^SSF.Scer\UAS under the control of Scer\GAL4^Bx-MS1096 in a smo³/+ background is suppressed by co-expression of smo^{Scer\UAS.cHa.T:Hsap\MYC}. The ectopic venation and costal overgrowth caused by expression of smo^{C.Scer\UAS.T:Hsap\MYC,T:Uuuu\Myr4} under the control of hemizygous Scer\GAL4^Bx-MS1096 in a smo³/+ background is enhanced by co-expression of fu^Scer\UAS.cAb, while the L3/4 narrowing is suppressed. (Hooper, 2003) Ommatidial clusters situated posterior to smo³; tkv⁸ double mutant clones differentiate properly, but over-rotate at a much higher frequency than those situated in other regions of the same eye disc, with many of them reaching 110^o-120^o. (Chou and Chien, 2002) smo³ homozygous clones generated in an en^Apa/+ background have no effect on cell fate or polarity in the tergites; smo³ mutant cells located posterior to the line of symmetry retain posterior orientation, while smo³ mutant cells located anterior to the line retain anterior orientation. The smo³ clones interdigitate freely with the surrounding cells. (Kopp and Duncan, 2002) Pka-C1^E95, sgg^M11 smo³ triple mutant clones in the wing disc exhibit some outgrowths from the notum but no significant wing pattern duplications. Wing discs with Pka-C1^E95, smo³ clones have large expansions affecting the anterior compartment of the wing disc. (Price and Kalderon, 2002) When Mad¹², smo³ double mutant somatic clones are made in the developing eye, a small but significant reduction in the percentage of mutant cells is seen compared to Mad¹² clones alone. (Rangarajan et al., 2001) smo³ Mad^1-2 double mutant clones in the eye never develop photoreceptors whether they lie at the margin or in the interior of the eye disc. Occasionally an ommatidium forms along the edge of the clone, such that some of the component photoreceptors lie within the clone. (Curtiss and Mlodzik, 2000) Clones in the posterior compartment of the wing disc which are expressing hh^αTub84B.PB in a smo³ background form wiggly borders with the surrounding cells. Clones in the anterior compartment of the wing disc which are expressing hh^αTub84B.PB in a smo³ background form significantly smoother borders with neighbouring wild-type cells. Unlike Df(2R)en^E single mutant clones, Df(2R)en^E smo³ double mutant clones of posterior origin in the wing disc invariably occupy only posterior territory and define straight borders to anterior cells at the normal position of the anterior/posterior (A/P) boundary. Df(2R)en^E smo³ double mutant clones of anterior origin also occupy posterior territory and define straight borders to anterior cells at the normal position of the A/P boundary. (Dahmann and Basler, 2000) Cells in tkv⁸ smo³ double mutant clones in the eye disc cannot differentiate as photoreceptors. (Greenwood and Struhl, 1999) Cells in the A compartment of the adult abdominal tergite which are simultaneously mutant for Df(2R)en^E and smo³ behave like smo³ clones: they transform a6, a5 and a4 into a3, and a1 into a2. (Lawrence et al., 1999) Clones mutant for smo³ and Df(2R)en^E induced in the posterior compartment of the abdominal tergite make a3 or a2 cuticle. The small number of clones produced after clone formation in the embryo tend to be elliptical with smooth boundaries. Clone induction in the larva produces mutant clones with less pigment and small bristles. These cells survive better than Df(2R)en^E clones, especially in the posterior part of the P compartment. The boundaries of these clones are smooth. Clones mutant for smo³ and Df(2R)en^E induced in the posterior compartment of the pleura form a3 or a2 cuticle embedded in the A compartments, having been ejected from the posterior compartment. (Lawrence et al., 1999) smo³; slmb¹/slmb² clones generate a supernumerary 'double-anterior' wing. (Jiang and Struhl, 1998) ptc^G12 homozygotes exhibit reduced number of denticle rows and each belt is rectangular shape. ptc^G12 smo³ double homozygotes raised at 25^oC show an essentially wild type phenotype. Scer\GAL4^h-1J3-mediated expression of hh^Scer\UAS.cIa at 18^oC does not modify the smo phenotype. Scer\GAL4^h-1J3-mediated expression of wg^Scer\UAS.cLa at 18^oC suppresses the smo denticle belt phenotype. (Quirk et al., 1997)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Rescued by	smo¹/smo³ is rescued by Scer\GAL4^prd.RG1/smo^{Scer\UAS.cHa.T:Hsap\MYC} (Hooper, 2003) smo³ is rescued by Scer\GAL4^prd.RG1/smo^{Scer\UAS.J.T:Avic\GFP} (Jia et al., 2003) smo³ is rescued by smo^+t6.2 (Chen and Struhl, 1998) smo³ is rescued by smo^{Scer\UAS.cvdHa}/Scer\GAL4^da.G32 (Zhu et al., 2003) smo³ is rescued by smo^{Scer\UAS.T:Avic\GFP}/Scer\GAL4^da.G32 (Zhu et al., 2003) smo³ is rescued by smo^{Scer\UAS.T:Hsap\MYC}/Scer\GAL4^prd.RG1 (Jia et al., 2003) smo³ is rescued by smo^{Scer\UAS.T:Zzzz\FLAG}/Scer\GAL4^arm.PS (Ingham et al., 2000)
Partially rescued by	smo³ is partially rescued by Scer\GAL4^prd.RG1/smo^{Scer\UAS.T:Zzzz\FLAG} (Nakano et al., 2004) smo³ is partially rescued by smo^{Scer\UAS.T:Zzzz\FLAG}/Scer\GAL4^h-1J3 (Ingham et al., 2000)
Not rescued by	smo¹/smo³ is not rescued by Scer\GAL4^prd.RG1/smo^{C.Scer\UAS.T:Hsap\MYC,T:Uuuu\Myr4} (Hooper, 2003) smo¹/smo³ is not rescued by Scer\GAL4^prd.RG1/smo^{N.Scer\UAS.T:Hsap\MYC} (Hooper, 2003) smo³ is not rescued by Scer\GAL4^prd.RG1/smo^{ΔCT.Scer\UAS.T:Avic\GFP} (Jia et al., 2003) smo³ is not rescued by smo^C155Y (Chen and Struhl, 1998)
Comments
Stocks ( 2 )
Bloomington	3277 smo³ b¹ pr¹/CyO
Kyoto	106954 smo³ b¹ pr¹/CyO
Notes on Origin
Discoverer

Comments
	The mutant phenotypes of smo² and smo³ are indistinguishable. (Strutt and Mlodzik, 1997)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 4 )
Reported As
Symbol Synonym	smo3 (Jia et al., 2010, Fan et al., 2012) smo³ (Vrailas et al., 2006, Wei et al., 2005, Molnar and de Celis, 2006, Jia et al., 2005, Kirilly et al., 2005, Sweeney et al., 2007, Firth and Baker, 2005, Casso et al., 2008, Cabernard and Affolter, 2005, Shi et al., 2011, Vrailas and Moses, 2006, Schlichting and Dahmann, 2008, Khaliullina et al., 2009, Baonza and Freeman, 2005, Corrigall et al., 2007, Jia et al., 2009, Nicholson et al., 2011, Schilling et al., 2011, Christiansen et al., 2012, Melicharek et al., 2008, Hartman et al., 2010) smo^Q14 (Thomas and Ingham, 2003, Ramirez-Weber et al., 2001, Jack and Myette, 1999, van den Heuvel and Ingham, 1996, Ma et al., 1996, Klingler and Gergen, 1993, Tearle and Nusslein-Volhard, 1987, ) smooth^Q14 (Parkhurst and Ish-Horowicz, 1991)
Name Synonym
Secondary FlyBase IDs
	FBal0032744
References ( 81 )

Generate a list of	All references relating to this allele ( 81 )

List References by type	Research paper ( 79 ) Stock list ( 1 ) Letter ( 1 )
Recent research papers ( 5 )
	Christiansen et al., 2012, Mech. Dev. 129(5-8): 98--108 Ligand-independent activation of the Hedgehog pathway displays non-cell autonomous proliferation during eye development in Drosophila. [FBrf0219046] Fan et al., 2012, Dev. Biol. 366(2): 172--184 Hh-induced Smoothened conformational switch is mediated by differential phosphorylation at its C-terminal tail in a dose- and position-dependent manner. [FBrf0218374] Nicholson et al., 2011, Development 138(2): 251--260 Notch-dependent expression of the archipelago ubiquitin ligase subunit in the Drosophila eye. [FBrf0212669] Schilling et al., 2011, PLoS Comput. Biol. 7(4): e1002025 Cell-sorting at the a/p boundary in the Drosophila wing primordium: a computational model to consolidate observed non-local effects of hh signaling. [FBrf0213439] Shi et al., 2011, Development 138(19): 4219--4231 The Hedgehog-induced Smoothened conformational switch assembles a signaling complex that activates Fused by promoting its dimerization and phosphorylation. [FBrf0216910] Show all research papers ...

Allele Dmel\smo3

Allele Dmel\smo³