Allele Dmel\mod(mdg4)^neo129

General Information
Symbol	Dmel\mod(mdg4)^neo129	Species	D. melanogaster
Name		FlyBase ID	FBal0003361
Feature type	allele	Associated gene	Dmel\mod(mdg4)
Also Known As	mod(mdg4)^E(var)3-93D, E(var)3-93D

Allele class	loss of function allele
Mutagen	P-element activity, Delta2-3
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	loss of function allele (Buchner et al., 2000)
Mutagen	Delta2-3 (Dorn et al., 1993) P-element activity (Buchner et al., 2000)
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 P{lacW} in the third intron at nucleotide 153,036 of AE003734. (Mongelard et al., 2002) Insertion into the third intron, which separates coding exon 2 from coding exon 3 of the mod(mdg4) transcription unit. The insertion lies 121 bp downstream from the 3' junction of coding exon 2. (Buchner et al., 2000) Insertion of a P-element into the third intron, 121bp downstream to the 5' splice junction. (Dorn et al., 1993)
Caused by insertion	P{hsneo.ry⁺}mod(mdg4)^neo129 (Dorn et al., 1993, Dorn et al., 1993, Buchner et al., 2000, Dorn, 1993.11.4, Sun et al., 2000, Soltani-Bejnood et al., 2007)
Cytology
Phenotypic Data
Phenotypic Class
	enhancer of variegation \| dominant, with In(1)w^m4 (Dorn et al., 1993, Dorn et al., 1993) enhancer of variegation \| dominant, with P{hsp26-pt-T}39C-42 (Sun et al., 2000) enhancer of variegation \| dominant, with P{hsp26-pt-T}118E-25-5 (Sun et al., 2000) female sterile (Dorn et al., 1993) female sterile (with mod(mdg4)⁰³) (Buchner et al., 2000) lethal (with mod(mdg4)⁰²) (Gabler et al., 2005) lethal (with mod(mdg4)⁰⁷) (Gabler et al., 2005) lethal \| cold sensitive (Dorn et al., 1993) lethal \| larval stage \| polyphasic \| recessive (Mongelard et al., 2002) lethal \| pupal stage (with mod(mdg4)^T16) (Chen and Corces, 2001) lethal \| pupal stage \| recessive (Gerasimova and Corces, 1998) lethal \| pupal stage \| recessive \| heat sensitive (Buchner et al., 2000) lethal \| recessive (Gabler et al., 2005) meiotic cell cycle defective \| male (with mod(mdg4)^Z3-3401) (Soltani-Bejnood et al., 2007) non-enhancer of variegation \| dominant, with P{hsp26-pt-T}39C-12 (Sun et al., 2000) non-enhancer of variegation \| dominant, with P{hsp26-pt-T}118E-3 (Sun et al., 2000) non-enhancer of variegation \| dominant, with P{hsp26-pt-T}118E-10 (Sun et al., 2000) non-enhancer of variegation \| dominant, with P{hsp26-pt-T}118E-19 (Sun et al., 2000) non-enhancer of variegation \| dominant, with P{hsp26-pt-T}CG2316^39C-34 (Sun et al., 2000) non-suppressor of variegation \| dominant, with T(2;3)lt^X13 (Weiler and Wakimoto, 2002) partially lethal - majority die (Soltani-Bejnood et al., 2007) partially lethal - majority die (with mod(mdg4)⁰³) (Buchner et al., 2000) suppressor of variegation \| dominant, with In(2L)lt^X2 (Weiler and Wakimoto, 2002) suppressor of variegation \| dominant, with In(2LR)lt^G10 (Weiler and Wakimoto, 2002)
Phenotype Manifest In
	ovariole (Buchner et al., 2000) ovary (Buchner et al., 2000)
Detailed Description
	Statement Reference mod(mdg4)^neo129/+ suppresses the variegation of lt due to In(2LR)lt^G10 (seen in In(2LR)lt^G10/Df(2L)lt10 animals derived from lt¹⁶/Df(2L)lt10 females), so that the percentage of pigmented Malpighian tubule cells is increased in the double mutant larvae. mod(mdg4)^neo129/+ suppresses the variegation of lt due to In(2L)lt^X2 (seen in In(2L)lt^X2/Df(2L)lt10 animals derived from lt¹⁶/Df(2L)lt10 females), so that the percentage of pigmented Malpighian tubule cells is increased in the double mutant larvae. mod(mdg4)^neo129/+ does not suppress the variegation of lt due to T(2;3)lt^X13 (seen in T(2;3)lt^X13/Df(2L)lt10 animals derived from lt¹⁶/Df(2L)lt10 females); there is no significant change in the percentage of pigmented Malpighian tubule. (Weiler and Wakimoto, 2002) Enhances the variegation in, i.e. reduces the eye pigmentation of, In(1)w^m4h, Su(var)2-1¹ males. Homozygous females obtained at 29^oC produce a small number of eggs that do not show signs of further development. Ovaries are of various size and ovarioles are fewer than normal and contain fewer egg chambers. (Buchner et al., 2000) Does not enhance transinactivation of bw caused by Tp(2;2)bw-DX7. (Sass and Henikoff, 1999) Some homozygous escapers develop to the adult stage. (Gerasimova and Corces, 1998) Enhances variegation of In(1)w^m4h. (Dorn et al., 1993) Strong dominant enhancer effect on w variegation in In(1)w^m4h. Homozygous males display a strong transformation of the fifth into the fourth abdominal segment, weak transformation is displayed by heterozygotes. (Dorn et al., 1993)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhanced by
Statement Reference In(1)w^m4, mod(mdg4)^neo129 has enhancer of variegation phenotype, enhanceable by trx^red-P6/trx[+] (Buchner et al., 2000)
Suppressed by
Statement Reference mod(mdg4)^neo129/mod(mdg4)⁰² has lethal phenotype, suppressible \| partially by Dvir\mod(mdg4)^t6.8 (Gabler et al., 2005) mod(mdg4)^neo129/mod(mdg4)⁰² has lethal phenotype, suppressible \| partially by Dvir\mod(mdg4)^t11.5 (Gabler et al., 2005) mod(mdg4)^neo129/mod(mdg4)^R32 has lethal phenotype, suppressible \| partially by Dvir\mod(mdg4)^t11.5 (Gabler et al., 2005) mod(mdg4)^neo129 has lethal \| recessive phenotype, suppressible \| partially by Dvir\mod(mdg4)^t11.5 (Gabler et al., 2005)
NOT suppressed by
Statement Reference mod(mdg4)⁰⁷/mod(mdg4)^neo129 has lethal phenotype, non-suppressible by Dvir\mod(mdg4)^t11.5 (Gabler et al., 2005)
Enhancer of
Statement Reference mod(mdg4)^neo129/mod(mdg4)[+] is an enhancer of enhancer of variegation phenotype of In(1)w^m4, vtd² (Buchner et al., 2000) mod(mdg4)^neo129/mod(mdg4)[+] is an enhancer of visible \| homeotic phenotype of E(var)3-4¹ (Buchner et al., 2000) mod(mdg4)^neo129/mod(mdg4)[+] is an enhancer of visible \| homeotic phenotype of trx^red-P6 (Buchner et al., 2000)
Suppressor of
Statement Reference mod(mdg4)^neo129/mod(mdg4)[+] is a suppressor of suppressor of variegation phenotype of In(1)w^m4h, Su(var)2-1¹ (Buchner et al., 2000)
NOT Suppressor of
Statement Reference mod(mdg4)^neo129/mod(mdg4)^ul is a non-suppressor of body color defective \| recessive phenotype of y² (Mongelard et al., 2002) mod(mdg4)^neo129/mod(mdg4)^ul is a non-suppressor of visible \| recessive phenotype of ct⁶ (Mongelard et al., 2002) mod(mdg4)^neo129/mod(mdg4)^ul is a non-suppressor of visible \| recessive phenotype of y² (Mongelard et al., 2002)
Phenotype Manifest In
Enhanced by
Statement Reference In(1)w^m4, mod(mdg4)^neo129 has pigment cell phenotype, enhanceable by trx^red-P6/trx[+] (Buchner et al., 2000)
Enhancer of
Statement Reference mod(mdg4)^neo129/mod(mdg4)[+] is an enhancer of adult abdominal segment 5 phenotype of E(var)3-4¹ (Buchner et al., 2000) mod(mdg4)^neo129/mod(mdg4)[+] is an enhancer of adult abdominal segment 5 phenotype of trx^red-P6 (Buchner et al., 2000) mod(mdg4)^neo129/mod(mdg4)[+] is an enhancer of pigment cell phenotype of In(1)w^m4, vtd² (Buchner et al., 2000)
NOT Suppressor of
Statement Reference mod(mdg4)^neo129/mod(mdg4)^ul is a non-suppressor of adult epidermis phenotype of y² (Mongelard et al., 2002) mod(mdg4)^neo129/mod(mdg4)^ul is a non-suppressor of wing phenotype of ct⁶ (Mongelard et al., 2002)
Other
Statement Reference In(1)w^m4h, Su(var)2-1¹, mod(mdg4)^neo129/mod(mdg4)[+] has pigment cell phenotype (Buchner et al., 2000)
Additional Comments
Genetic Interactions
Statement Reference Enhances the homeotic transformation of A5 to A4 of trx^red-P6 and E(var)3-4¹. (Buchner et al., 2000)
Xenogenetic Interactions
Statement Reference One copy of Dvir\mod(mdg4)^t11.5 can rescue the lethality of mod(mdg4)^neo129 homozygotes (92% of the expected progeny survive). One copy of Dvir\mod(mdg4)^t11.5 can rescue the lethality of mod(mdg4)⁰²/mod(mdg4)^neo129 transheterozygotes, with the degree of rescue depending on the P{Dvir\mod(mdg4)^t11.5} line used (95.1% and full viability has been seen). One copy of Dvir\mod(mdg4)^t11.5 can rescue the lethality of mod(mdg4)^R32/mod(mdg4)^neo129 transheterozygotes. Rescued males show greater viability (63.2%) compared to rescued females (40.5%). One copy of Dvir\mod(mdg4)^t11.5 does not suppresses the lethality of mod(mdg4)⁰⁷/mod(mdg4)^neo129 transheterozygotes. One copy of Dvir\mod(mdg4)^t6.8 partially rescues the lethality of mod(mdg4)⁰²/mod(mdg4)^neo129 transheterozygotes (34% viability is seen). (Gabler et al., 2005)
Complementation & Rescue Data
Complements	mod(mdg4)^T6 (Mongelard et al., 2002) mod(mdg4)^ul (Mongelard et al., 2002)
Partially complements	mod(mdg4)⁰³ (Buchner et al., 2000) mod(mdg4)^Z3-3401 (Soltani-Bejnood et al., 2007)
Fails to complement	mod(mdg4)⁰² (Buchner et al., 2000) mod(mdg4)⁰⁴ (Buchner et al., 2000) mod(mdg4)⁰⁶ (Buchner et al., 2000) mod(mdg4)⁰⁷ (Buchner et al., 2000) mod(mdg4)⁰³⁸⁵² (Mongelard et al., 2002) mod(mdg4)^bpd1 (Gorczyca et al., 1999) mod(mdg4)^bpdEX1 (Gorczyca et al., 1999) mod(mdg4)^L3101 (Mongelard et al., 2002) mod(mdg4)^R32 (Buchner et al., 2000)
Partially rescued by	mod(mdg4)^neo129 is partially rescued by mod(mdg4)^+t7.5 (Buchner et al., 2000, Soltani-Bejnood et al., 2007, Soltani-Bejnood et al., 2007) mod(mdg4)^neo129 is partially rescued by mod(mdg4)^t7.5 (Dorn et al., 2001)
Comments	The semi-lethal phenotype of mod(mdg4)^neo129 is fully complemented by mod(mdg4)^Z3-3401, but mod(mdg4)^neo129 fails to complement mod(mdg4)^Z3-3401 with respect to X-Y chromosome nondisjunction in male meiosis - transheterozygotes show a high level of X-Y nondisjunction (41.2%). The lethal phenotype of mod(mdg4)^neo129 is fully complemented by mod(mdg4)^Z3-5578 and mod(mdg4)^Z3-3298, and mod(mdg4)^neo129 partially or fully complements these alleles with respect to X-Y chromosome nondisjunction in male meiosis - the transheterozygotes show only a low level of X-Y nondisjunction (frequency is indicated in parentheses after each allele); mod(mdg4)^Z3-5578 (4.46%) and mod(mdg4)^Z3-3298 (7.64%). (Soltani-Bejnood et al., 2007) Member of Group I complementation group. Complementation statement based on transheterozygous phenotype in terms of suppression or non-suppression of y² and ct⁶ visible phenotypes. (Mongelard et al., 2002) Complementation statements based on viability. (Buchner et al., 2000)
Stocks ( 0 )

Notes on Origin
Discoverer

Comments
	Alleles of mod(mdg4) show an allelic series for their effect on In(1)w^m4h variegation. From strongest to weakest: mod(mdg4)^R32 = mod(mdg4)⁰⁴ = mod(mdg4)⁰⁶ = mod(mdg4)⁰⁷ = mod(mdg4)^142Δ32 > mod(mdg4)^neo129 = mod(mdg4)⁰² = mod(mdg4)⁰³ > mod(mdg4)⁰⁵. (Buchner et al., 2000) P-element has been precisely excised generating wild type revertants. (Dorn et al., 1993) Strong enhancer. High spontaneous background rate of E(var) mutations in these experiments allows for the possibility that the P insertion is not the cause of the E(var) phenotype. (Dorn et al., 1993)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 8 )
Reported As
Symbol Synonym	E(var)3-93D (Sun et al., 2000, Sass and Henikoff, 1999) E(var)3-93D-neo129 (Gorczyca et al., 1999) E(var)93D^neo129 E(var)93D^neo (Buchner et al., 1995) E-var(3)3⁰¹ (Dorn et al., 1993) mod(mdg4)^E(var)3-93D (Mongelard et al., 2002, Chen and Corces, 2001, Gerasimova and Corces, 1998) mod(mdg4)^neo129 neo129 (Soltani-Bejnood et al., 2007)
Name Synonym
Secondary FlyBase IDs

References ( 15 )
Research paper	Soltani-Bejnood et al., 2007, Genetics 176(1): 161--180 Role of the mod(mdg4) common region in homolog segregation in Drosophila male meiosis. [FBrf0201774] Gabler et al., 2005, Genetics 169(2): 723--736 Trans-splicing of the mod(mdg4) complex locus is conserved between the distantly related species Drosophila melanogaster and D. virilis. [FBrf0188488] Mongelard et al., 2002, Genetics 160(4): 1481--1487 Trans-splicing as a novel mechanism to explain interallelic complementation in Drosophila. [FBrf0147059] Weiler and Wakimoto, 2002, Mol. Genet. Genomics 266(6): 922--932 Suppression of heterochromatic gene variegation can be used to distinguish and characterize E(var) genes potentially important for chromosome structure in Drosophila melanogaster. [FBrf0145133] Chen and Corces, 2001, Genetics 159(4): 1649--1658 The gypsy insulator of Drosophila affects chromatin structure in a directional manner. [FBrf0141553] Dorn et al., 2001, Proc. Natl. Acad. Sci. U.S.A. 98(17): 9724--9729 Transgene analysis proves mRNA trans-splicing at the complex mod(mdg4) locus in Drosophila. [FBrf0137352] Buchner et al., 2000, Genetics 155(1): 141--157 Genetic and molecular complexity of the position effect variegation modifier mod(mdg4) in Drosophila. [FBrf0127028] Sun et al., 2000, Proc. Natl. Acad. Sci. U.S.A. 97(10): 5340--5345 The fourth chromosome of Drosophila melanogaster: interspersed euchromatic and heterochromatic domains. [FBrf0127342] Gorczyca et al., 1999, J. Neurobiol. 39(3): 447--460 The gene mod(mdg4) affects synapse specificity and structure in Drosophila. [FBrf0108771] Sass and Henikoff, 1999, Genetics 152(2): 595--604 Pairing-dependent mislocalization of a Drosophila brown gene reporter to a heterochromatic environment. [FBrf0109049] Gerasimova and Corces, 1998, Cell 92(4): 511--521 Polycomb and trithorax group proteins mediate the function of a chromatin insulator. [FBrf0100803] Dorn et al., 1993, Genetics 133(2): 279--290 P transposon-induced dominant enhancer mutations of position-effect variegation in Drosophila melanogaster. [FBrf0058558] Dorn et al., 1993, Proc. Natl. Acad. Sci. U.S.A. 90(23): 11376--11380 The enhancer of position-effect variegation of Drosophila, E(var)3-93D, codes for a chromatin protein containing a conserved domain common to several transcriptional regulators. [FBrf0064436]
Abstract	Buchner et al., 1995, Europ. Dros. Res. Conf. 14: 322 E(var)3-93D - Molecular characterization and expression during development. [FBrf0083057]
DNA/RNA sequence record	Dorn, 1993.11.4, GenBank/EMBL/DDBJ: X75498 D. melanogaster mRNA for proteinA. [FBrf0114628]

Allele Dmel\mod(mdg4)neo129

Allele Dmel\mod(mdg4)^neo129