Allele Dmel\ptc^tuf-1

General Information
Symbol	Dmel\ptc^tuf-1	Species	D. melanogaster
Name		FlyBase ID	FBal0028491
Feature type	allele	Associated gene	Dmel\ptc
Also Known As	ptc^tuf1, ptc^tuf

Allele class	hypomorphic allele - genetic evidence
Mutagen	spontaneous, ethyl methanesulfonate
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	hypomorphic allele - genetic evidence (Mullor and Guerrero, 2000)
Mutagen	ethyl methanesulfonate (Lindsley and Zimm, 1992) spontaneous (Phillips et al., 1990, Capdevila et al., 1994)
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 Insertion of 7kb of DNA in the 5' regulatory region, located between -50 and -100bp upstream of the transcription initiation site. Insertion of about 7kb in the 5' regulatory region between -50 and -100bp upstream of the transcription initiation site. (Capdevila et al., 1994)
Cytology
Phenotypic Data
Phenotypic Class
	increased cell death (with ptc¹³) (Thomas and Ingham, 2003) increased cell death (with ptc¹⁶) (Thomas and Ingham, 2003) lethal \| recessive (Lindsley and Zimm, 1992) viable (Capdevila et al., 1994, Phillips et al., 1990, Mullor and Guerrero, 2000) visible (Lindsley and Zimm, 1992, Sturtevant, 1948) visible (with ptc¹³) (Strutt et al., 2001) visible (with ptc³⁴) (Strutt et al., 2001) visible (with ptc^S2) (Strutt et al., 2001)
Phenotype Manifest In
	1st posterior cell (Strutt et al., 2001) adult head (Sturtevant, 1948) antenna (with ptc⁹) (Thomas and Ingham, 2003) antenna (with ptc¹⁶) (Thomas and Ingham, 2003) antenna (with ptc¹⁷) (Thomas and Ingham, 2003) antenna (with ptc³⁷) (Thomas and Ingham, 2003) antenna (with ptc⁴⁷) (Thomas and Ingham, 2003) antenna (with ptc^G12) (Thomas and Ingham, 2003) antennal disc (with ptc¹³) (Thomas and Ingham, 2003) costal cell (Capdevila et al., 1994, Strutt et al., 2001) double row \| ectopic (with ptc¹³) (Strutt et al., 2001) double row \| ectopic (with ptc³⁴) (Strutt et al., 2001) eye (with ptc⁹) (Thomas and Ingham, 2003) eye (with ptc¹³) (Thomas and Ingham, 2003) eye (with ptc¹⁴) (Thomas and Ingham, 2003) eye (with ptc¹⁵) (Thomas and Ingham, 2003) eye (with ptc¹⁶) (Thomas and Ingham, 2003) eye (with ptc¹⁷) (Thomas and Ingham, 2003) eye (with ptc³⁴) (Thomas and Ingham, 2003) eye (with ptc³⁷) (Thomas and Ingham, 2003) eye (with ptc⁴⁷) (Thomas and Ingham, 2003) eye (with ptc^G12) (Thomas and Ingham, 2003) eye (with ptc^H84) (Thomas and Ingham, 2003) eye (with ptc^S2) (Thomas and Ingham, 2003) eye disc (with ptc¹³) (Thomas and Ingham, 2003) eye disc (with ptc³⁷) (Thomas and Ingham, 2003) frons (with ptc⁹) (Thomas and Ingham, 2003) frons (with ptc¹⁴) (Thomas and Ingham, 2003) frons (with ptc¹⁵) (Thomas and Ingham, 2003) frons (with ptc¹⁶) (Thomas and Ingham, 2003) frons (with ptc¹⁷) (Thomas and Ingham, 2003) frons (with ptc³⁴) (Thomas and Ingham, 2003) frons (with ptc³⁷) (Thomas and Ingham, 2003) frons (with ptc⁴⁷) (Thomas and Ingham, 2003) frons (with ptc^G12) (Thomas and Ingham, 2003) frons (with ptc^H84) (Thomas and Ingham, 2003) frons (with ptc^S2) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc⁹) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc¹⁴) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc¹⁵) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc¹⁶) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc¹⁷) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc³⁴) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc³⁷) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc⁴⁷) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc^G12) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc^H84) (Thomas and Ingham, 2003) fronto-orbital plate (with ptc^S2) (Thomas and Ingham, 2003) head (with ptc^tuf-2) (Lindsley and Zimm, 1992) head bristle (with ptc⁹) (Thomas and Ingham, 2003) head bristle (with ptc¹³) (Thomas and Ingham, 2003) head bristle (with ptc¹⁴) (Thomas and Ingham, 2003) head bristle (with ptc¹⁵) (Thomas and Ingham, 2003) head bristle (with ptc¹⁶) (Thomas and Ingham, 2003) head bristle (with ptc¹⁷) (Thomas and Ingham, 2003) head bristle (with ptc³⁴) (Thomas and Ingham, 2003) head bristle (with ptc³⁷) (Thomas and Ingham, 2003) head bristle (with ptc⁴⁷) (Thomas and Ingham, 2003) head bristle (with ptc^559.1) (Thomas and Ingham, 2003) head bristle (with ptc^G12) (Thomas and Ingham, 2003) head bristle (with ptc^H84) (Thomas and Ingham, 2003) head bristle (with ptc^S2) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc⁹) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc¹³) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc¹⁴) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc¹⁵) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc¹⁶) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc¹⁷) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc³⁴) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc³⁷) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc⁴⁷) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc^559.1) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc^G12) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc^H84) (Thomas and Ingham, 2003) head bristle \| ectopic (with ptc^S2) (Thomas and Ingham, 2003) head capsule & cuticle \| ectopic (with ptc⁹) (Thomas and Ingham, 2003) head capsule & cuticle \| ectopic (with ptc¹³) (Thomas and Ingham, 2003) head capsule & cuticle \| ectopic (with ptc¹⁶) (Thomas and Ingham, 2003) head capsule & cuticle \| ectopic (with ptc¹⁷) (Thomas and Ingham, 2003) head capsule & cuticle \| ectopic (with ptc³⁴) (Thomas and Ingham, 2003) head capsule & cuticle \| ectopic (with ptc³⁷) (Thomas and Ingham, 2003) head capsule & cuticle \| ectopic (with ptc⁴⁷) (Thomas and Ingham, 2003) head capsule & cuticle \| ectopic (with ptc^G12) (Thomas and Ingham, 2003) head capsule & cuticle \| ectopic (with ptc^S2) (Thomas and Ingham, 2003) macrochaeta (Capdevila et al., 1994) ocellar triangle (with ptc⁹) (Thomas and Ingham, 2003) ocellar triangle (with ptc¹⁴) (Thomas and Ingham, 2003) ocellar triangle (with ptc¹⁵) (Thomas and Ingham, 2003) ocellar triangle (with ptc¹⁶) (Thomas and Ingham, 2003) ocellar triangle (with ptc¹⁷) (Thomas and Ingham, 2003) ocellar triangle (with ptc³⁴) (Thomas and Ingham, 2003) ocellar triangle (with ptc³⁷) (Thomas and Ingham, 2003) ocellar triangle (with ptc⁴⁷) (Thomas and Ingham, 2003) ocellar triangle (with ptc^G12) (Thomas and Ingham, 2003) ocellar triangle (with ptc^H84) (Thomas and Ingham, 2003) ocellar triangle (with ptc^S2) (Thomas and Ingham, 2003) ocellus \| ectopic (with ptc⁹) (Thomas and Ingham, 2003) ocellus \| ectopic (with ptc¹⁶) (Thomas and Ingham, 2003) ocellus \| ectopic (with ptc¹⁷) (Thomas and Ingham, 2003) ocellus \| ectopic (with ptc³⁷) (Thomas and Ingham, 2003) ocellus \| ectopic (with ptc⁴⁷) (Thomas and Ingham, 2003) ommatidium (with ptc⁹) (Thomas and Ingham, 2003) ommatidium (with ptc¹⁴) (Thomas and Ingham, 2003) ommatidium (with ptc¹⁵) (Thomas and Ingham, 2003) ommatidium (with ptc¹⁶) (Thomas and Ingham, 2003) ommatidium (with ptc¹⁷) (Thomas and Ingham, 2003) ommatidium (with ptc³⁷) (Thomas and Ingham, 2003) ommatidium (with ptc⁴⁷) (Thomas and Ingham, 2003) ommatidium (with ptc^G12) (Thomas and Ingham, 2003) scutellar bristle (Capdevila et al., 1994) scutellum (Mullor et al., 1997) triple row (with ptc¹³) (Strutt et al., 2001) triple row (with ptc³⁴) (Strutt et al., 2001) wing (Phillips et al., 1990) wing (with ptc¹³) (Strutt et al., 2001) wing (with ptc³⁴) (Strutt et al., 2001) wing (with ptc^Conf-1) (Mullor and Guerrero, 2000) wing (with ptc^S2) (Strutt et al., 2001) wing (with ptc^tuf-2) (Lindsley and Zimm, 1992) wing \| anterior (Sturtevant, 1948) wing \| anterior compartment (Capdevila et al., 1994) wing disc (Canamasas et al., 2003) wing margin (with ptc¹³) (Strutt et al., 2001) wing margin (with ptc³⁴) (Strutt et al., 2001) wing margin (with ptc^S2) (Strutt et al., 2001) wing vein (Capdevila et al., 1994, Phillips et al., 1990) wing vein L3 (with ptc^Conf-1) (Mullor and Guerrero, 2000) wing vein L4 (with ptc^Conf-1) (Mullor and Guerrero, 2000)
Detailed Description
	Statement Reference Mutants show overgrowth of the central part of the anterior region of the wing imaginal disc. (Canamasas et al., 2003) All flies carrying ptc^tuf-1 in trans to ptc³⁷, ptc¹⁷, ptc¹⁶, ptc⁴⁷ or ptc⁹ have severely reduced, rough eyes, a highly enlarged head vertex (ocellar triangle, fronto-orbital plate and frons), outgrowths of head cuticle and large numbers of missing, misplaced or ectopic head bristles. In addition around 9-50% of these flies have missing antennae and 10-56% have ectopic ocelli. ptc^G12/ptc^tuf-1 heads are much like those just described except that head vertex enlargement is not so severe and ectopic ocelli are not seen. ptc^tuf-1/ptc^H84 flies have mildly reduced eyes but a highly enlarged head vertex. 90% of these flies have outgrowths of head cuticle and large numbers of missing, misplaced or ectopic head bristles with the remaining 10% having no outgrowths and one or 2 displaced or missing bristles. Flies carrying ptc^tuf-1 in trans to ptc¹⁵ or ptc¹⁴ have mildly reduced, rough eyes and an enlarged head vertex. Around 30% also have one or two missing or misplaced head bristles. ptc^tuf-1/ptc^559.1 have generally wild-type looking heads, with only 5% showing missing or misplaced head bristles. ptc³⁴/ptc^tuf-1 and ptc^S2/ptc^tuf-1 flies have enlarged eyes and enlarged head vertex (ocellar triangle, fronto-orbital plate and frons). Other head defects in these flies vary from outgrowths of head cuticle plus large numbers of missing, misplaced or ectopic head bristles to one or 2 misplaced or missing head bristles. ptc¹³/ptc^tuf-1 have enlarged eyes. The head vertex in these flies is of normal size, but all have outgrowths of head cuticle plus large numbers of missing, misplaced or ectopic head bristles. The zone of cell cycle arrest corresponding to the morphogenetic furrow (MF) is not significantly increased in late 3rd instar ptc¹³/ptc^tuf-1 or ptc³⁷/ptc^tuf-1 eye discs compared to wild-type. However, its width is uneven along the length of the furrow, being particularly narrow at the equator and wider along the margins. The distribution of dividing cells either side of the MF is disorganized: dividing cells anterior and posterior to the furrow are closer together near the equator of the disc than at the poles. Compared with wild type, there is an approximately 2 fold increase in cell division ahead of the furrow in the mutants. This is particularly apparent in the ptc¹³/ptc^tuf-1 animals and thus could partially provide an explanation for the large-eye phenotype. In the eye discs of late third instar ptc¹⁶/ptc^tuf-1 larvae, apoptosis is increased in a broad swath of cells situated behind the furrow. ptc¹³/ptc^tuf-1 mutant eye discs also show increased cell death despite the resulting enlarged eye although it is not as extensive as in ptc¹⁶/ptc^tuf-1. Occasionally a vast amount of cell death is seen in the antennal disc of ptc¹³/ptc^tuf-1 animals. (Thomas and Ingham, 2003) homozygous and hemizygous mutants display distortion in the costal cell and an increase in the distance between the L3 and L4 wing veins. Mutants transheterozygous for ptc^tuf-1 and ptc^S2, ptc¹³ or ptc³⁴ show normal separation of wing veins L3 and L4, but display dramatic shape changes and abnormalities in the anterior margin. With ptc¹³ the wing becomes almost circular and the triple row bristles are replaced by double row bristles. With ptc^S2, mirror duplications of the proximal half of the anterior wing are seen. (Strutt et al., 2001) In ptc^tuf-1/ptc^Conf-1 flies the development of the central region of the wing is affected, vein 4 partially develops, and vein 3 is thickened. (Mullor and Guerrero, 2000) ptc^tuf-1/ptc¹⁶ flies have extra bristles in the scutellum. (Mullor et al., 1997) Coordinate mutant. (Sturtevant and Bier, 1995) Heterozygotes with ptc¹⁶ and ptc⁹ display a variable phenotype including overgrowth of the anterior compartment of the wing, loss of costal structures and wing vein defects and loss of scutellar bristles. P{HSptc.S} construct can rescue the mutant phenotype. Viable. ptc^tuf-1/ptc⁹ and ptc^tuf-1/ptc¹⁶ heterozygotes display severe outgrowth of the anterior compartment of the wing disc, loss of costal structures, duplications or plexations of veins 1 and 2, increase of the distance between veins 3 and 4 and increase in the number of scutellar bristles. (Capdevila et al., 1994) Wings of ptc^tuf-1/ptc^tuf-2 grossly foreshortened and shaped like a ping-pong paddle; these flies also show duplications and triplications of anterior wing structures, lack costal bristles and have more head abnormalities than ptc^tuf-1 homozygotes. Extreme allele. (Lindsley and Zimm, 1992) ptc^tuf-1 homozygotes display a variable wing phenotype, ranging from wild-type to loss of costal bristles and moderate duplication of the base of vein 1 and associated triple row bristles. ptc^tuf-1 in combination with lethal ptc alleles cause more extensive deletions of the costa and duplications of veins 1 and 2. Triple row bristles often show a change in orientation between the sites at which the duplicated veins touch the margin. (Phillips et al., 1990) Flies have small tufts of hair between the eye and antenna, and show basal twinning of the anterior half of the wing. (Sturtevant, 1948) viable
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Phenotype Manifest In
Enhanced by
Statement Reference ptc^tuf-1 has phenotype, enhanceable by ci^D (Sanchez-Herrero et al., 1996)
Suppressed by
Statement Reference ptc^Conf-1/ptc^tuf-1 has wing phenotype, suppressible by Scer\GAL4^hh-Gal4/hh^Scer\UAS.cCa (Mullor and Guerrero, 2000) ptc^Conf-1/ptc^tuf-1 has wing vein L3 phenotype, suppressible by Scer\GAL4^hh-Gal4/hh^Scer\UAS.cCa (Mullor and Guerrero, 2000) ptc^Conf-1/ptc^tuf-1 has wing vein L4 phenotype, suppressible by Scer\GAL4^hh-Gal4/hh^Scer\UAS.cCa (Mullor and Guerrero, 2000)
Suppressor of
Statement Reference ptc¹³/ptc^tuf-1 is a suppressor of eye phenotype of hh^bar3 (Thomas and Ingham, 2003) ptc¹⁴/ptc^tuf-1 is a suppressor \| partially of eye phenotype of hh^bar3 (Thomas and Ingham, 2003) ptc¹⁶/ptc^tuf-1 is a suppressor \| partially of eye phenotype of hh^bar3 (Thomas and Ingham, 2003) ptc³⁴/ptc^tuf-1 is a suppressor \| partially of eye phenotype of hh^bar3 (Thomas and Ingham, 2003) ptc³⁷/ptc^tuf-1 is a suppressor \| partially of eye phenotype of hh^bar3 (Thomas and Ingham, 2003) ptc⁴⁷/ptc^tuf-1 is a suppressor \| partially of eye phenotype of hh^bar3 (Thomas and Ingham, 2003)
Additional Comments
Genetic Interactions
Statement Reference l(2)tid¹ ptc^tuf-1 double homozygous larvae develop tumourous imaginal discs and die at the larval/pupal transition, just as l(2)tid¹ homozygotes do. (Canamasas et al., 2003) The phenotype is significantly increased by ci^D. (Sanchez-Herrero et al., 1996)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Partially complements	ptc^Conf-1 (Mullor and Guerrero, 2000)
Rescued by	ptc^tuf-1 is rescued by ptc^hs.PSa (Capdevila et al., 1994)
Comments
Stocks ( 4 )
Bloomington	429 ptc^tuf-1 ltd¹ 239 w¹¹¹⁸; Bl¹ stw³ ap^blt ptc^tuf-1 sp¹/SM6a
Kyoto	105908 ptc^tuf-1 ltd¹ 105808 w¹¹¹⁸; Bl¹ stw³ ap^blt ptc^tuf-1 sp¹/SM6a
Notes on Origin
Discoverer

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 9 )
Reported As
Symbol Synonym	ptc^tf (Sturtevant and Bier, 1995) ptc^tuf1 (Thomas and Ingham, 2003, Mullor and Guerrero, 2000, Mullor et al., 1997, Sanchez-Herrero et al., 1996) ptc^tuf-1 ptc^Tuf1 (Ayers et al., 2009) ptc^tuf (Strutt et al., 2001, Johnson et al., 1995, Phillips et al., 1990) ptctuf1 (Thomas and Ingham, 2003) tuf (Sturtevant, 1948) tuf¹ (Capdevila et al., 1994, ) tuf² (Lindsley and Zimm, 1992)
Name Synonym
Secondary FlyBase IDs

References ( 14 )
Research paper	Ayers et al., 2009, Dev. Biol. 329(2): 280--293 Tow (Target of Wingless), a novel repressor of the Hedgehog pathway in Drosophila. [FBrf0208057] Canamasas et al., 2003, J. Biol. Chem. 278(33): 30952--30960 Understanding human cancer using Drosophila: Tid47, a cytosolic product of the DnaJ-like tumour suppressor gene l(2)Tid, is a novel molecular partner of patched related to skin cancer. [FBrf0173155] Thomas and Ingham, 2003, Genetics 165(4): 1915--1928 Hedgehog signaling in the Drosophila eye and head: an analysis of the effects of different patched trans-heterozygotes. [FBrf0167651] Strutt et al., 2001, Curr. Biol. 11(8): 608--613 Mutations in the sterol-sensing domain of Patched suggest a role for vesicular trafficking in Smoothened regulation. [FBrf0136834] Mullor and Guerrero, 2000, Dev. Biol. 228(2): 211--224 A gain-of-function mutant of patched dissects different responses to the Hedgehog gradient. [FBrf0132285] Mullor et al., 1997, Development 124(6): 1227--1237 Hedgehog activity, independent of decapentaplegic, participates in wing disc patterning. [FBrf0093168] Sanchez-Herrero et al., 1996, Mech. Dev. 55(2): 159--170 The fu gene discriminates between pathways to control dpp expression in Drosophila imaginal discs. [FBrf0087686] Johnson et al., 1995, Development 121(12): 4161--4170 patched overexpression alters wing disc size and pattern: transcriptional and post-transcriptional effects on hedgehog targets. [FBrf0085299] Sturtevant and Bier, 1995, Development 121(3): 785--801 Analysis of the genetic hierarchy guiding wing vein development in Drosophila. [FBrf0076146] Capdevila et al., 1994, EMBO J. 13(1): 71--82 The Drosophila segment polarity gene patched interacts with decapentaplegic in wing development. [FBrf0068440] Phillips et al., 1990, Development 110: 105--114 The Drosophila segment polarity gene patched is involved in a position-signalling mechanism in imaginal discs. [FBrf0051546] Sturtevant, 1948, D. I. S. 22: 55--56 [New mutants report.] [FBrf0063893]
Abstract	Thomas and Ingham, 2003, Europ. Dros. Res. Conf. 18: A46 An Analysis Of Eye And Head Phenotypes In ptc Transheterozygotes Of Drosophila. [FBrf0161668]
Book	Lindsley and Zimm, 1992, The Genome of Drosophila melanogaster. The Genome of Drosophila melanogaster. [FBrf0066905]

Allele Dmel\ptctuf-1

Allele Dmel\ptc^tuf-1