Aberration Dmel\In(1)w^m4

General Information
Symbol	Dmel\In(1)w^m4	Species	D. melanogaster
Name	Inversion (1) white-mottled	FlyBase ID	FBab0004257
Feature type	chromosomal_inversion	Created / Updated	2006-08-22/2006-08-22

Formalized genetic data	l(1)2Ea << bk1 << w << CR << bk2
Sequence coordinates
Deleted segment
Duplicated segment
Computed Breakpoints include	3C2;h28
Breakpoints Inherited
Nature of the Aberration
Cytological Order
Progenitor
Mutagen	X ray
Class of aberration (relative to progenitor)	chromosomal_inversion (Tartof et al., 1984, Gersh, 1967, Slobodyanyuk and Serov, 1983, The Moscow Regional Drosophila melanogaster Stock Center, Dubna)
Breakpoints	3C2;20B (Gersh, 1967) 3C1-3C2;20A (Slobodyanyuk and Serov, 1983) 3C1-3C2;h28 (Tartof et al., 1984) 3C2-3C3;20B (The Moscow Regional Drosophila melanogaster Stock Center, Dubna)
Causes alleles
Carries alleles
Transposon Insertions
Genetic mapping information
Comments	Left (3C1-2) breakpoint at -24.5 kb in the restriction map of the w locus (0 point at site of the copia insertion in w^a); right (20F) breakpoint near 5' end of a Type I mobile element (Tartof et al., 1984). Left breakpoint less than 3kb distal to the w-a copia insertion (Pirrotta et al., 1983). Breakpoint(s) molecularly mapped (Pirrotta et al., 1983)
Comments on Cytology
	Limits of break 1 from polytene analysis (FBrf0018623) Limits of break 2 from polytene analysis (FBrf0040503) Proximal breakpoint defines the distal boundary of the bb locus. (Appels and Hilliker, 1982) Distal inversion breakpoint is ~20kb from the 3' end of the w locus, the proximal inversion breakpoint is within centric heterochromatin distal to bb. Heterochromatic breakpoint is flanked by a Type I mobile element (FBrf0040503). (Judd, 1995) Proximal breakpoint is approximate. The distal breakpoint is to the left of w and the proximal breakpoint is to the left of bb. (Gersh, 1967)
Molecularly Mapped Breakpoints

Sequence Crossreferences
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name
Gene Deletion & Duplication Data
Genes Deleted / Disrupted
Complementation Data
Molecular Data
Genes NOT Deleted / Disrupted
Complementation Data
Molecular Data
Genes Duplicated
Complementation Data
Molecular Data
Genes NOT Duplicated
Complementation Data
Molecular Data
Related Comments

Phenotypic Data
In combination with other aberrations
NOT in combination with other aberrations	In(1)w^m4 flies show position effect variegation at the w locus, at 23^oC the phenotype varies from a sectored red and white eye to a red eye peppered with small flecks of red and brown pigmented ommatidia. (Locke et al., 1988) Rpd3^15-1 enhances the variegation of w in In(1)w^m4 flies and in 5-10% of the eyes patches of ommatidia show disorganisation characteristic of the rst mutation (enhancement of variegation allows the silencing to spread past the w locus to inactivate rst in a fraction of the cells). (De Rubertis et al., 1996) Tn10\tetR^{ey.3.5.T:Hsim\VP16}-mediated expression of Hsap\HMGIY^{MATH20.Tn10\tetO} in In(1)w^m4 flies results in significant derepression of the w gene and a general loss of the variegating phenotype. Tetracycline treatment inhibits Hsap\HMGIY^{MATH20.Tn10\tetO}-induced suppression of PEV. Tn10\tetR^{ey.3.5.T:Hsim\VP16}-mediated expression of Hsap\HMGIY^{MATH11.Tn10\tetO} in In(1)w^m4 flies results in no significant derepression of the w gene, no loss of the variegating phenotype. (Girard et al., 1998) Wow^γb, Wow^γe and Wow^hd1 act as suppressors of variegation of w in In(1)w^m4. (Birchler et al., 1994) z⁺ In(1)w^m4 flies reared at 25^oC have dark red/brown mottled eyes. z¹ In(1)w^m4 males and females have fewer red/brown patches on a lighter background, paired copies of In(1)w^m4 are not repressed to z eye colour (lemon yellow). Repression of In(1)w^m4 by z¹ cannot be restored by Su(var)205. z^v77h In(1)w^m4 females and males have almost bleached white eye colour with a few scattered red facets, Su(var)205 only moderately suppresses PEV. (Judd, 1995) Carnitine compounds (L-Carnitine, L-Acetylcarnitine and L-Propionylcarnitine) show a significant suppression on w variegation. Butyrate gives a weaker suppression. In males the suppression effect of the compounds was seen at all concentrations, but in females the effect is weak and is only seen with high concentrations. (Fanti et al., 1994) Enhances the expression of Pgd^A in larvae and adults. (Slobodyanyuk and Serov, 1983) Eyes have a "sectored" phenotype - ommatidia are either fully pigmented or not pigmented at all. (Tartof et al., 1984) Males and homozygous females are viable and fertile. (Lefevre and Wilkins, 1966) The position effect variegation at the w locus caused by In(1)w^m4 is enhanced by Su(var)3-7^+t6.5. (Cleard et al., 1997) The position effect variegation caused by In(1)w^m4 is dominantly suppressed by Su(z)12¹. (Birve and Rasmuson-Lestander, 1994) Variegated eye phenotype is enhanced in the presence of mod(mdg4)^ul, mod(mdg4)^u2 and mod(mdg4)^B2, eyes appear yellow with orange spots. In the presence of su(Hw)^MC the phenotype is no longer enhanced and the eye phenotype returns to wild type. These results indicate that the variegating phenotype is caused by the su(Hw) protein. (Gerasimova et al., 1995)
Position Effect Variegation Data
Genes subject to PEV	w (Sinclair et al., 1998, Sass and Henikoff, 1998, Girard et al., 1998, Zorin et al., 1999, Talbert and Henikoff, 2000, Mottus et al., 2000, Shareef et al., 2001, Lefevre and Wilkins, 1966, Gersh, 1967, Tartof et al., 1984, Locke et al., 1988, Sinclair et al., 1989, Tartof and Bremer, 1990, Spofford and DeSalle, 1991, Judd, 1995, De Rubertis et al., 1996, Lloyd et al., 1997, Eberl et al., 1997, Spofford, 1965, Dorer and Henikoff, 1997)
Stocks ( 23 )
Bloomington	807 In(1)w^m4 490 In(1)w^m4; Df(2L)E110/CyO 1712 In(1)w^m4; Df(2L)2802/CyO 6904 In(1)w^m4; Df(2L)J106/CyO 9576 In(1)w^m4; hup¹/CyO 994 In(1)w^m4; T(2;4)Su(var)2/Cy¹ 1304 In(1)w^m4, In(1)AB, y²/C(1)M3 3805 Df(YS)bb-Su(var)/In(1)w^m4, w¹
Kyoto	101140 In(1)w^m4 106083 In(1)w^m4 106466 In(1)w^m4; Df(2L)2802/CyO 108906 In(1)w^m4; Df(2L)E110/CyO
	More stocks available...
Notes on Origin
Discoverer	Muller, 1929.

Balancer / Genotype Variants of the Aberration

Separable Components

Other Comments
	A strongly variegating line has been designated In(1)w^m4h. X ray-induced revertants of In(1)w^m4 may or may not carry a segment of flanking heterochromatin: variegation is not controlled from immediately adjacent heterochromatic sequences at the euchromatin/ heterochromatin border but from a site more internal to the heterochromatin domain. (Tartof et al., 1989) w transvection is eliminated by PEV. (Judd, 1995) The presence of a variegating chromosome and a modifier chromosome in the same parental genome can alter the amount the amount of variegation formed in the progeny. The genomic imprinting is not determined by the parental origin of the variegating chromosome but is instead determined by the genetic background the variegating chromosome is subjected to during gametogenesis. (Bishop and Jackson, 1996)
Synonyms & Secondary IDs ( 12 )
Reported As
Symbol Synonym	In(1)^wm4 (Gonzy et al., 2002) In(1)w^m4 (Lerach et al., 2006, Nakayama et al., 2007, Schwendemann et al., 2008) In(1)wm4 (Boivin and Dura, 1998) In(1)w-m4 m⁴ (Slobodyanyuk and Serov, 1983) w^4-2880 (Donaldson et al., 2002) w^m4 (Haley et al., 2005, Badugu et al., 2003, Malmanche and Clark, 2003, Schotta et al., 2002, Tulin et al., 2002, Granok et al., 2001, Reeves, 2001, Janssen et al., 2000, Shareef et al., 2001, Henikoff and Vermaak, 2000, Green and Piergentili, 2000, Buchner et al., 2000, Lefevre and Wilkins, 1966, Hart and Laemmli, 1998, Lefevre, 1968, Sass and Henikoff, 1998, Sinclair et al., 1998, Henikoff, 1990, Spradling and Karpen, 1990, Bezborodova et al., 1997, Larsson and Rasmuson-Lestander, 1997, The Moscow Regional Drosophila melanogaster Stock Center, Dubna, Tartof and Bremer, 1990, Cleard et al., 1997, Henikoff, 1996, Park and Yamamoto, 1995, Moehrle and Paro, 1994, Lloyd et al., 1997, Gdula et al., 1996, Shaffer et al., 1993, Tartof et al., 1984, Birchler et al., 1994, Sinclair et al., 1992, Weiler, 2001, Mason and Konev, 2001, Mason et al., 2002, Weiler, 2004, Bao et al., 2006, Lerach et al., 2006, McHugh et al., 2004, Maines et al., 2007, Rudolph et al., 2007, Eggert et al., 2004) white^m4 (Bornemann et al., 2002) wm4 (Eggert et al., 2003)
Name Synonym	Inversion (1) white-mottled white^mottled4 (Gvozdev et al., 2000) white mottled 4 (McHugh et al., 2004)
Secondary FlyBase IDs
	FBal0018268
References ( 141 )

Generate a list of	All references relating to this aberration ( 141 )

List References by type	Research paper ( 104 ) Review ( 17 ) Personal communication to FlyBase ( 1 ) Abstract ( 16 ) FlyBase analysis ( 1 ) Stock list ( 1 ) Book ( 1 )
Recent research papers ( 6 )
	Schwendemann et al., 2008, Proc. Natl. Acad. Sci. USA 105(1): 204--209 Hip, an HP1-interacting protein, is a haplo- and triplo-suppressor of position effect variegation. [FBrf0203015] Maines et al., 2007, Development 134(8): 1471--1479 Stonewalling Drosophila stem cell differentiation by epigenetic controls. [FBrf0200664] Nakayama et al., 2007, Genes Dev. 21(5): 552--561 Drosophila GAGA factor directs histone H3.3 replacement that prevents the heterochromatin spreading. [FBrf0195269] Rudolph et al., 2007, Molec. Cell 26(1): 103--115 Heterochromatin Formation in Drosophila Is Initiated through Active Removal of H3K4 Methylation by the LSD1 Homolog SU(VAR)3-3. [FBrf0204799] Blattes et al., 2006, EMBO J. 25(11): 2397--2408 Displacement of D1, HP1 and topoisomerase II from satellite heterochromatin by a specific polyamide. [FBrf0194408] Lerach et al., 2006, Genetics 173(4): 2403--2406 Loss-of-function alleles of the JIL-1 kinase are strong suppressors of position effect variegation of the wm4 allele in Drosophila. [FBrf0192731] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2006 Show reviews ...

Aberration Dmel\In(1)wm4

Aberration Dmel\In(1)w^m4