Allele Dmel\Pc¹⁵

General Information
Symbol	Dmel\Pc¹⁵	Species	D. melanogaster
Name		FlyBase ID	FBal0013565
Feature type	allele	Associated gene	Dmel\Pc
Also Known As	Pc^XT109

Allele class	amorphic allele - genetic evidence
Mutagen	X ray
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	amorphic allele - genetic evidence (Simon et al., 1992)
Mutagen	X ray (Tearle and Nusslein-Volhard, 1987, Strutt and Paro, 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
Nature of the lesion	Statement Reference
Cytology
Phenotypic Data
Phenotypic Class
	decreased cell number \| somatic clone (Bello et al., 2007) mitotic cell cycle defective \| embryonic stage (O'Dor et al., 2006) mitotic cell cycle defective \| somatic clone (Bello et al., 2007) neuroanatomy defective \| somatic clone (Wang et al., 2006, Parrish et al., 2007) suppressor of variegation \| dominant, with P{UAS-lacZ.Abd-B.5F24}25,2a (Grienenberger et al., 2002) tumorigenic \| somatic clone (Classen et al., 2009, Gutiérrez et al., 2012) visible \| dominant (Strutt et al., 1997, Culi et al., 2006, Anderson et al., 2011)
Phenotype Manifest In
	anaphase \| embryonic stage (O'Dor et al., 2006) dendrite & dorsal multidendritic neuron ddaC \| somatic clone (Parrish et al., 2007) dendrite \| somatic clone (Wang et al., 2006) eye disc \| somatic clone (Classen et al., 2009, Gutiérrez et al., 2012) gonadal sheath proper primordium (with Pc³) (DeFalco et al., 2004) haltere \| ectopic (Culi et al., 2006) larval abdominal segment 4 (Grienenberger et al., 2002) larval abdominal segment 5 \| ectopic (Grienenberger et al., 2002) larval head (Muller et al., 1995) larval segment (Muller et al., 1995) mesothoracic leg (Strutt et al., 1997, Lindsley and Zimm, 1992, Anderson et al., 2011) mesothoracic leg & sex comb \| ectopic (Strutt et al., 1997) metathoracic leg (Lindsley and Zimm, 1992, Grienenberger et al., 2002, Anderson et al., 2011) neuroblast \| somatic clone (Bello et al., 2007) sex comb (with Df(3R)ry27) (Wang and Brock, 2003) sex comb (with Df(3R)ry619) (Wang and Brock, 2003) sex comb \| ectopic (Strutt et al., 1997, Grienenberger et al., 2002, Anderson et al., 2011, Wen et al., 2012) telophase \| embryonic stage (O'Dor et al., 2006) wing (Culi et al., 2006)
Detailed Description
	Statement Reference Eye discs composed mainly of homozygous cells have a tumour phenotype. (Gutiérrez et al., 2012) Approximately 90% of Pc[15] heterozygotes have extra sex combs. (Wen et al., 2012) 82% of heterozygous males have ectopic sex combs on either the second or third leg. (Anderson et al., 2011) Eye discs composed predominantly of homozygous cells (generated using the FLP/cell-lethal method) are overgrown. The mutant tissue shows defects in terminal differentiation and has defective epithelial organisation. (Classen et al., 2009) Homozygous neuroblast clones in the third larval instar central brain and ventral ganglion are dramatically reduced in size compared to control clones and show a dramatic reduction in mitotic activity. Homozygous neuroblast clones examined at the late third larval stage generally still contain neuroblasts (88% of clones) if examined 24 hours after clone induction, but neuroblasts are generally absent in the clones (89% of clones) 72 hours after clone induction. 59% of the mutant clones contain neuroblasts 48 hours after clone induction. (Bello et al., 2007) Single cell class IV dendrite arborisation (da) neuron clones that are homozygous for Pc[15] show a range of dendritic defects in the third instar larva. Homozygous class IV ddaC clones show a significant reduction in dendrite branching and total dendritic length compared to controls. Analysis of mutant clones from 72 to 96 hours after egg laying (AEL) shows that mutant clones show extensive growth of major dendritic branches and some terminal branches. However, roughly a quarter of the terminal dendrites are lost in the mutant clones during this time period. Fewer terminal branching/branch growth events are seen in the mutant clones than the control clones. Axons of mutant single cell ddaC neuron clones occupy a similar fascicular position in the ventral nerve cord and show very similar arborisation patterns as wild-type controls. Axon terminals of mutant v'ada or vdab clones are also indistinguishable from wild type. (Parrish et al., 2007) Heterozygotes show a partial transformation of wing to haltere. (Culi et al., 2006) Pc¹⁵ embryos from hetrozygous mothers show segregation defects in anaphase and telophase as judged by the presence of chromatin bridges. Most of these chromatin bridges are probably resolved because large irregularly-shaped nuclei and polyploid nuclei are only seen at a low penetrance. Pc¹⁵ embryos also show a higher level of "nuclear fallout", a process that removes nuclei with abnormal mitoses, than wild-type embryos. Fallout nuclei tend to be observed in pairs or clusters and have a late telophase appearance. There is no evidence of metaphase defects in these embryos. (O'Dor et al., 2006) Pc¹⁵ mushroom body clones show over-elaborated dendrites, but do not show similar axonal defects. (Wang et al., 2006) Ectopic male-specific somatic gonadal precursor (msSGP) cells are seen in both male and female Pc³/Pc¹⁵ embryos before gonad coalescence. Normal and ectopic non-sexually dimorphic somatic gonadal precursor cell (SGP) specification is also seen in these mutant embryos. At later stage in mutant male embryos, a large number of ectopic msSGPs associate with SGPs and germ cells to form a gonad. However, in most mutant female embryos msSGPs are no longer seen at later stages. (DeFalco et al., 2004) Pc¹⁵/Df(3R)ry27 mutant flies exhibit a slightly higher than wild-type average number of legs with sex combs, with a statistically insignificant average of 2.1 legs with sex combs, compared to 2.0 in Pc¹⁵/+ heterozygotes. Pc¹⁵/Df(3R)ry619 mutant flies exhibit a higher than wild-type average number of legs with sex combs, with a statistically significant average of 2.2 legs with sex combs, compared to 2.0 in Pc¹⁵/Tp(3;3)MRS heterozygotes. (Wang and Brock, 2003) Pc¹⁵/+ animals exhibit extra sex combs on the metathoracic leg, they also have a mild transformation of the fourth abdominal segment into the fifth - patches of pigmentation are seen in the anterior part of A4. (Grienenberger et al., 2002) Heterozygous males have ectopic sex combs on the second leg. (Strutt et al., 1997) Segments of homozygous larvae are transformed so they mostly represent the eighth abdominal segment and the larvae show severe head defects. (Muller et al., 1995) Partial transformations of second and third legs into first legs in Pc/+ heterozygotes. (Lindsley and Zimm, 1992) The loss of the zygotic Pc causes a severe transformation of most segments towards parasegment 8 (Lewis, Nature 276: 565-570, Duncan, Developmental Order, pp 533--554, New York). (Simon et al., 1992)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhanced by
Statement Reference Pc¹⁵ has visible \| dominant phenotype, enhanceable by jing[+]/jing^22F3 (Culi et al., 2006)
NOT Enhanced by
Statement Reference Pc¹⁵ has neuroanatomy defective \| somatic clone phenotype, non-enhanceable by esc²¹/esc[+] (Parrish et al., 2007)
Suppressed by
Statement Reference Pc¹⁵ has tumorigenic \| somatic clone phenotype, suppressible \| somatic clone by dome^{ΔCYT.Scer\UAS}/Scer\GAL4^Act.PU (Classen et al., 2009) Pc¹⁵ has visible \| dominant phenotype, suppressible \| partially by Caf1[+]/Caf1^long (Anderson et al., 2011) Pc¹⁵ has visible \| dominant phenotype, suppressible \| partially by Caf1^short/Caf1[+] (Anderson et al., 2011)
NOT suppressed by
Statement Reference Pc¹⁵ has neuroanatomy defective \| somatic clone phenotype, non-suppressible by esc²¹/esc[+] (Parrish et al., 2007)
Enhancer of
Statement Reference Pc[+]/Pc¹⁵ is an enhancer of visible \| dominant phenotype of gcm^Pyx (Popkova et al., 2012)
Suppressor of
Statement Reference Pc[+]/Pc¹⁵ is a suppressor \| partially of eye color defective phenotype of w^PD (Classen et al., 2009) Pc[+]/Pc¹⁵ is a suppressor \| partially of eye color defective phenotype of w^PRECycA (Martinez et al., 2006) Pc¹⁵ is a suppressor of eye color defective phenotype of w^Fab-7.5F24 (Cavalli, 1999) Pc¹⁵ is a suppressor of visible phenotype of Scer\GAL4^lz-gal4, sens^Scer\UAS.cNa (Anderson et al., 2011)
Phenotype Manifest In
Enhanced by
Statement Reference Pc¹⁵ has haltere \| ectopic phenotype, enhanceable by jing[+]/jing^22F3 (Culi et al., 2006) Pc¹⁵ has larval abdominal segment 4 phenotype, enhanceable by chm¹⁴ (Grienenberger et al., 2002) Pc¹⁵ has larval abdominal segment 4 phenotype, enhanceable by chm¹⁴/Abd-B^Mcp-B116 (Grienenberger et al., 2002) Pc¹⁵ has larval abdominal segment 5 \| ectopic phenotype, enhanceable by chm¹⁴ (Grienenberger et al., 2002) Pc¹⁵ has larval abdominal segment 5 \| ectopic phenotype, enhanceable by chm¹⁴/Abd-B^Mcp-B116 (Grienenberger et al., 2002) Pc¹⁵ has mesothoracic leg phenotype, enhanceable by Trl^13C (Strutt et al., 1997) Pc¹⁵ has mesothoracic leg phenotype, enhanceable by Trl⁶² (Strutt et al., 1997) Pc¹⁵ has metathoracic leg phenotype, enhanceable by chm¹⁴ (Grienenberger et al., 2002) Pc¹⁵ has phenotype, enhanceable by Scm^ET50 (Bornemann et al., 1998) Pc¹⁵ has phenotype, enhanceable by Scm^R5-13 (Bornemann et al., 1998) Pc¹⁵ has phenotype, enhanceable by Scm^Suz302 (Bornemann et al., 1998) Pc¹⁵ has sex comb \| ectopic & mesothoracic leg phenotype, enhanceable by Trl[+]/Trl^13C (Strutt et al., 1997) Pc¹⁵ has sex comb \| ectopic & mesothoracic leg phenotype, enhanceable by Trl[+]/Trl⁶² (Strutt et al., 1997) Pc¹⁵ has sex comb \| ectopic phenotype, enhanceable by chm¹⁴ (Grienenberger et al., 2002) Pc¹⁵ has wing phenotype, enhanceable by jing[+]/jing^22F3 (Culi et al., 2006)
NOT Enhanced by
Statement Reference Pc¹⁵ has dendrite & dorsal multidendritic neuron ddaC \| somatic clone phenotype, non-enhanceable by esc²¹/esc[+] (Parrish et al., 2007) Pc¹⁵ has sex comb \| ectopic phenotype, non-enhanceable by Caf1[+]/Caf1^p55-1 (Wen et al., 2012) Pc¹⁵ has sex comb \| ectopic phenotype, non-enhanceable by Caf1^p55-2/Caf1[+] (Wen et al., 2012)
Suppressed by
Statement Reference Pc¹⁵ has eye disc \| somatic clone phenotype, suppressible \| somatic clone by dome^{ΔCYT.Scer\UAS}/Scer\GAL4^Act.PU (Classen et al., 2009) Pc¹⁵ has larval head phenotype, suppressible \| partially by Mmus\Cbx2^arm.PM (Muller et al., 1995) Pc¹⁵ has larval segment phenotype, suppressible \| partially by Mmus\Cbx2^arm.PM (Muller et al., 1995) Pc¹⁵ has mesothoracic leg phenotype, suppressible \| partially by Caf1[+]/Caf1^long (Anderson et al., 2011) Pc¹⁵ has mesothoracic leg phenotype, suppressible \| partially by Caf1^short/Caf1[+] (Anderson et al., 2011) Pc¹⁵ has metathoracic leg phenotype, suppressible \| partially by Caf1[+]/Caf1^long (Anderson et al., 2011) Pc¹⁵ has metathoracic leg phenotype, suppressible \| partially by Caf1^short/Caf1[+] (Anderson et al., 2011) Pc¹⁵ has neuroblast \| somatic clone phenotype, suppressible by Scer\GAL4^αTub84B.PL/BacA\p35^Scer\UAS.cHa (Bello et al., 2007) Pc¹⁵ has sex comb \| ectopic phenotype, suppressible \| partially by Caf1[+]/Caf1^long (Anderson et al., 2011) Pc¹⁵ has sex comb \| ectopic phenotype, suppressible \| partially by Caf1^short/Caf1[+] (Anderson et al., 2011)
NOT suppressed by
Statement Reference Pc¹⁵ has dendrite & dorsal multidendritic neuron ddaC \| somatic clone phenotype, non-suppressible by esc²¹/esc[+] (Parrish et al., 2007) Pc¹⁵ has sex comb \| ectopic phenotype, non-suppressible by Caf1[+]/Caf1^p55-1 (Wen et al., 2012) Pc¹⁵ has sex comb \| ectopic phenotype, non-suppressible by Caf1^p55-2/Caf1[+] (Wen et al., 2012)
Enhancer of
Statement Reference chm¹⁴/Pc¹⁵ is an enhancer of larval abdominal segment 4 phenotype of Abd-B^Mcp-B116 (Grienenberger et al., 2002) chm¹⁴/Pc¹⁵ is an enhancer of larval abdominal segment 5 \| ectopic phenotype of Abd-B^Mcp-B116 (Grienenberger et al., 2002) Pc[+]/Pc¹⁵ is an enhancer of chaeta \| supernumerary phenotype of gcm^Pyx (Popkova et al., 2012) Pc¹⁵/Abd-B^Mcp-B116 is an enhancer of larval abdominal segment 4 phenotype of chm¹⁴ (Grienenberger et al., 2002) Pc¹⁵/Abd-B^Mcp-B116 is an enhancer of larval abdominal segment 5 \| ectopic phenotype of chm¹⁴ (Grienenberger et al., 2002)
Suppressor of
Statement Reference Pc[+]/Pc¹⁵ is a suppressor \| partially of eye phenotype of w^PD (Classen et al., 2009) Pc[+]/Pc¹⁵ is a suppressor \| partially of pigment cell phenotype of w^PRECycA (Martinez et al., 2006) Pc¹⁵ is a suppressor of eye phenotype of Scer\GAL4^lz-gal4, sens^Scer\UAS.cNa (Anderson et al., 2011) Pc¹⁵ is a suppressor of pigment cell phenotype of w^Fab-7.5F24 (Cavalli, 1999)
Additional Comments
Genetic Interactions
Statement Reference Caf1[p55-1]/+ does not affect the ectopic sex comb phenotype seen in Pc[15] heterozygotes. Caf1[p55-2]/+ does not affect the ectopic sex comb phenotype seen in Pc[15] heterozygotes. (Wen et al., 2012) The disorganised eye phenotype caused by expression of sens[Scer\UAS.cNa] under the control of Scer\GAL4[lz-gal4] is suppressed by Pc[15]. Only 12.3% of Pc[15]/Caf1[short] and 30% of Pc[13]/Caf1[long] double heterozygous males have ectopic sex combs on either the second or third leg. (Anderson et al., 2011) Expression of dome[ΔCYT.Scer\UAS] under the control of Scer\GAL4[Act.PU] reduces the overgrowth of Pc[15] mutant eye discs. (Classen et al., 2009) The addition of esc[21]/+ does not alter the dendritic defects seen in single cell class IV dendrite arborisation (da) neuron clones that are homozygous for Pc[15]. (Parrish et al., 2007) The partial transformation of wing to haltere seen in Pc¹⁵ heterozygotes is enhanced by jing^22F3/+. (Culi et al., 2006) Mutants heterozygous for lolal^k02512, lolal^k11212, or lolal^k07907, when also heterozygous for Pc¹⁵, exhibit a contrabithorax phenotype, which is a wing-to-haltere transformation due to ectopic expression of Ubx in the posterior part of the wing disc, often resulting in curvature of the wing. This phenotype is further enhanced in specific lolal heteroallelic viable combinations, such as lolal^k02512/lolal^311a, lolal^k11212/lolal^311a, and lolal^k07907/lolal^311a, when also heterozygous for Pc¹⁵. (Faucheux et al., 2003) Mutant heterozygotes show a weak suppression of the eye phenotype seen in P{UAS-lacZ.Abd-B.5F24} leading to darker eyes. In Pc¹⁵/+, w^Fab-7.5F24, Scer\GAL4^hs.PB flies, an embryonic heat shock pulse leads to eyes darker than with w^Fab-7.5F24 alone. (Cavalli, 1999) The appearance of extra sex combs in Pc¹⁵/+ males is enhanced if the flies are also heterozygous for either Trl⁶² or Trl^13C. (Strutt et al., 1997)
Xenogenetic Interactions
Statement Reference Mutant phenotype can be rescued in the presence of Mmus\Cbx2^arm.PM. Larvae were not rescued to adults with Mmus\Cbx2^arm.PM. Clones of Pc¹⁵ adult cells can be rescued by one copy of Mmus\Cbx2^arm.PM but to an intermediate phenotype. (Muller et al., 1995)
Complementation & Rescue Data
Comments	Mutant phenotype can be rescued in the presence of Pc^arm.PM Clones of Pc¹⁵ adult cells can be completely rescued with one copy of Pc^arm.PM. (Muller et al., 1995)
Stocks ( 1 )
Bloomington	24468 w^*; P{hsp70-CD2.J}76 kni^ri-1 Pc¹⁵ P{FRT(w^hs)}2A/TM6B, Tb¹
Notes on Origin
Discoverer

Comments
	Ectopic abd-A is seen throughout the A-P axis by 9 hours. Ectopic Abd-B follows the same pattern but the anterior spread is delayed. Patchy Abd-B is seen in the epidermal cells of the head region. (Simon et al., 1992)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 4 )
Reported As
Symbol Synonym	Pc¹⁵ (Wang and Brock, 2003, de Ayala Alonso et al., 2007, Anderson et al., 2011, Zhai et al., 2009, Wen et al., 2012, Popkova et al., 2012) Pc^XT109 (Martinez et al., 2006, Zirin and Mann, 2004, Erhardt et al., 2003, DeFalco et al., 2004, Grienenberger et al., 2002, Grienenberger et al., 2002, Beuchle et al., 2001, Cavalli, 1999, Fritsch et al., 1999, Kehle et al., 1998, Bornemann et al., 1998, Strutt and Paro, 1997, Strutt et al., 1997, Muller et al., 1995, Tearle and Nusslein-Volhard, 1987, O'Dor et al., 2006, Wang et al., 2006, Culi et al., 2006, Bello et al., 2007, Chen et al., 2005, Parrish et al., 2007, Le Bras and Van Doren, 2006, Estella et al., 2008, Gutiérrez et al., 2012) Pc^XT (Atchison et al., 2003) XT109 (Tariq et al., 2009)
Name Synonym
Secondary FlyBase IDs

References ( 38 )

Generate a list of	All references relating to this allele ( 38 )

List References by type	Research paper ( 33 ) Supplementary material ( 2 ) Review ( 1 ) Stock list ( 1 ) Book ( 1 )
Recent research papers ( 4 )
	Gutiérrez et al., 2012, Development 139(1): 117--127 The role of the histone H2A ubiquitinase Sce in Polycomb repression. [FBrf0216868] Popkova et al., 2012, PLoS Genet. 8(12): e1003159 Polycomb controls gliogenesis by regulating the transient expression of the gcm/glide fate determinant. [FBrf0220515] Wen et al., 2012, Dev. Dyn. 241(3): 455--464 The biological function of the WD40 repeat-containing protein p55/Caf1 in Drosophila. [FBrf0217472] Anderson et al., 2011, Development 138(10): 1957--1966 The enhancer of trithorax and polycomb gene Caf1/p55 is essential for cell survival and patterning in Drosophila development. [FBrf0213580] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2011 Show reviews ...

Allele Dmel\Pc15

Allele Dmel\Pc¹⁵