Allele Dmel\mr²

General Information
Symbol	Dmel\mr²	Species	D. melanogaster
Name		FlyBase ID	FBal0012433
Feature type	allele	Associated gene	Dmel\mr

Map ( GBrowse )
Allele class
Mutagen	spontaneous
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class
Mutagen	spontaneous
Mutations Mapped to the Genome

Type Location Additional Notes References point mutation 2R:19,862,669..19,862,669 evidence=experimental pr_change=W739R\|mr-PA reported_pr_change=W739R na_change=T19862669C comment=Same mutation as mr¹. Site of nucleotide substitution in mutant inferred by FlyBase based on reported amino acid change. (Kashevsky et al., 2002)
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: W739R. (Kashevsky et al., 2002)
Cytology
Phenotypic Data
Phenotypic Class
	female sterile (Nagy et al., 2012) female sterile \| recessive (Reed and Orr-Weaver, 1997) sterile (with mr¹) (Kashevsky et al., 2002)
Phenotype Manifest In
	egg (Reed and Orr-Weaver, 1997) embryo (Reed and Orr-Weaver, 1997) eye (Reed and Orr-Weaver, 1997) follicle cell (Reed and Orr-Weaver, 1997) mitotic cell cycle \| embryonic stage (Reed and Orr-Weaver, 1997) nuclear chromosome \| maternal effect (with mr¹) (Kashevsky et al., 2002) nurse cell (with mr¹) (Kashevsky et al., 2002) nurse cell & nuclear chromosome (Reed and Orr-Weaver, 1997) nurse cell & nucleus (Reed and Orr-Weaver, 1997) nurse cell ring canal (Reed and Orr-Weaver, 1997) polar body nucleus (Reed and Orr-Weaver, 1997) spindle \| embryonic stage (Reed and Orr-Weaver, 1997) spindle \| maternal effect (with mr¹) (Kashevsky et al., 2002) tergite (Reed and Orr-Weaver, 1997)
Detailed Description
	Statement Reference mr[1]/mr[2] females show no apparent defect in germline stem cell maintenance. (Chen et al., 2009) The nurse cells revert to mitosis at stage 5 in mr¹/mr² egg chambers. Nurse cells in later mr¹/mr² egg chambers often become pycnotic. Embryos derived from mr¹/mr² egg chambers which complete oogenesis complete meiosis, however, the zygotic nuclei and frequently the polar bodies, are arrested on metaphase spindles that are anastral or have broad poles. The chromosomes are hypercondensed in these embryos. (Kashevsky et al., 2002) Semi-lethal when hemizygous, with most hemizygotes dying as pharate adults. Escapers have etched tergites and very rough eyes. Nurse cells in homozygous ovaries initiate endo cycles, but later egg chambers contain condensed chromosomes in place of polyploid nurse cell nuclei. These condensed chromosomes are associated with large multipolar spindles. Ring canals are seen to degenerate shortly after the inappropriate spindles are seen. The nurse cell defects are not visible until stage 4 of egg chamber development. The follicle cell epithelium sometimes forms a double layer of cells. Homozygous or mr¹/mr² females sometimes lay eggs (which do not hatch). Fertilised eggs show an early metaphase arrest phenotype, containing usually between 6 and 10 metaphase nuclei. The spindles are broader than wild-type both at the metaphase plate and at the spindle poles. Metaphase spindles are found at the position where the polar bodies are normally located. (Reed and Orr-Weaver, 1997) Less extreme than mr. Nearly wild type at 19^oC. female-sterile Oogenesis normal through stage 4; then compound nurse cell chromosomes fall apart and degenerate. Karyosome of oocyte also disappears. Oogenesis does not proceed beyond sixth stage (King, 1964). RK2 at 25^oC or above.
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Suppressor of
Statement Reference mr[+]/mr² is a suppressor of increased cell death phenotype of shtd¹ (Tanaka-Matakatsu et al., 2007)
Other
Statement Reference aret^PA/aret^Z2286, mr¹/mr² has mitotic cell cycle defective \| oogenesis stage S1 phenotype (Sugimura and Lilly, 2006) aret^PA/aret^Z2286, mr¹/mr² has mitotic cell cycle defective \| oogenesis stage S2 phenotype (Sugimura and Lilly, 2006) lmgA⁰³⁴²⁴, mr² has lethal \| prepupal stage P4(ii) phenotype (Nagy et al., 2012) lmgA⁰³⁴²⁴, mr² has mitotic cell cycle defective \| third instar larval stage phenotype (Nagy et al., 2012) mr², vih^KG02013 has lethal \| third instar larval stage phenotype (Nagy et al., 2012) mr², vih^S110501 has lethal \| third instar larval stage phenotype (Nagy et al., 2012)
Phenotype Manifest In
Enhanced by
Statement Reference mr² has phenotype, enhanceable by e(mr)¹ (Reed, 1992)
Enhancer of
Statement Reference mr¹/mr² is an enhancer of female germline stem cell phenotype of eff^mer1/eff⁸ (Chen et al., 2009)
Suppressor of
Statement Reference mr[+]/mr² is a suppressor of ommatidium phenotype of shtd¹ (Tanaka-Matakatsu et al., 2007)
Other
Statement Reference aret^PA/aret^Z2286, mr¹/mr² has germline cell \| female \| oogenesis stage S1 phenotype (Sugimura and Lilly, 2006) aret^PA/aret^Z2286, mr¹/mr² has germline cell \| female \| oogenesis stage S2 phenotype (Sugimura and Lilly, 2006) e(mr)¹, mr¹/mr² has antennal lobe phenotype (Reed, 1992) lmgA⁰³⁴²⁴, mr² has embryonic/larval brain \| third instar larval stage phenotype (Nagy et al., 2012)
Additional Comments
Genetic Interactions
Statement Reference lmgA[03424] mr[2] double mutants die at prepupal stage P4(ii). The mitotic index in third larval instar brains is increased compared to wild type and 31.8 +/- 3.9 % of the cells are polyploid. Third larval instar brains and wing imaginal discs show increased levels of apoptosis compared to wild type. (Nagy et al., 2012) mr[1]/mr[2] enhances the loss of germline stem cells that is seen in eff[8]/eff[mer1] females, such that the rate of germline stem cell loss is more rapid in the double mutant females. (Chen et al., 2009) In aret^PA/aret^Z2286 ; mr¹/mr² double mutant adult female, germ cells show a high frequency of metaphase arrest beginning during stage 1-2 of oogenesis. (Sugimura and Lilly, 2006)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Rescued by	mr¹/mr² is rescued by Scer\GAL4^Act5C.PI/Scer\GAL4^Act5C.PI/mr^{Scer\UAS.P\T.cKa} (Kashevsky et al., 2002) mr¹/mr² is rescued by Scer\GAL4^{nos.UTR.T:Hsim\VP16}/mr^{Scer\UAS.P\T.cKa} (Kashevsky et al., 2002)
Comments
Stocks ( 3 )
Bloomington	4535 e(mr)¹; mr²/SM6a 257 chl¹ l(2)bw¹ bw^2b mr²/SM5
Kyoto	107785 e(mr)¹; mr²/SM6a
Notes on Origin
Discoverer	Bridges, 24th Nov. 1925.

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 2 )
Reported As
Symbol Synonym	mr² (Nagy et al., 2012, Sugimura and Lilly, 2006, Tanaka-Matakatsu et al., 2007, Chen et al., 2009)
Name Synonym	morula² (Narbonne-Reveau et al., 2008)
Secondary FlyBase IDs

References ( 8 )
Research paper	Nagy et al., 2012, Cell Div. 7(1): 9 lemmingA encodes the Apc11 subunit of the APC/C in Drosophila melanogaster that forms a ternary complex with the E2-C type ubiquitin conjugating enzyme, Vihar and Morula/Apc2. [FBrf0218534] Chen et al., 2009, Development 136(24): 4133--4142 Effete-mediated degradation of Cyclin A is essential for the maintenance of germline stem cells in Drosophila. [FBrf0209346] Narbonne-Reveau et al., 2008, Development 135(8): 1451--1461 APC/CFzr/Cdh1 promotes cell cycle progression during the Drosophila endocycle. [FBrf0204408] Tanaka-Matakatsu et al., 2007, Dev. Biol. 309(2): 222--235 Mutation of the Apcl homologue shattered disrupts normal eye development by disrupting G1 cell cycle arrest and progression through mitosis. [FBrf0201230] Sugimura and Lilly, 2006, Dev. Cell 10(1): 127--135 Bruno inhibits the expression of mitotic cyclins during the prophase I meiotic arrest of Drosophila oocytes. [FBrf0190220] Kashevsky et al., 2002, Proc. Natl. Acad. Sci. U.S.A. 99(17): 11217--11222 The anaphase promoting complex/cyclosome is required during development for modified cell cycles. [FBrf0152304] Reed and Orr-Weaver, 1997, Development 124(18): 3543--3553 The Drosophila gene morula inhibits mitotic functions in the endo cell cycle and the mitotic cell cycle. [FBrf0098873]
Thesis	Reed, 1992, Ph.D. Thesis, University of Cambridge, England: 292pp The genetic analysis of endoreduplication in Drosophila melanogaster. [FBrf0064297]

Allele Dmel\mr2

Allele Dmel\mr²