Allele Dmel\Mer³

General Information
Symbol	Dmel\Mer³	Species	D. melanogaster
Name		FlyBase ID	FBal0061732
Feature type	allele	Associated gene	Dmel\Mer

Map ( GBrowse )
Allele class
Mutagen	ethyl methanesulfonate
Recent Updates
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FB2012_01
FB2011_10
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Nature of the Allele
Allele class
Mutagen	ethyl methanesulfonate (Fehon et al., 1997)
Mutations Mapped to the Genome

Type Location Additional Notes References point mutation X:19,586,460..19,586,460 evidence=experimental reported_pr_change=M177I comment=Site of nucleotide substitution in mutant inferred by FlyBase based on reported amino acid change. Nucleotide substitution not specified (could be G to A, G to T, or G to C) pr_change=M177I\|Mer-PA (LaJeunesse et al., 1998)
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: M177I. (LaJeunesse et al., 1998)
Cytology
Phenotypic Data
Phenotypic Class
	cytokinesis defective (Dorogova et al., 2008) decreased cell size \| recessive (McCartney et al., 2000) lethal \| male \| partially \| recessive (LaJeunesse et al., 2001) meiotic cell cycle defective (Dorogova et al., 2008) sterile (MacDougall et al., 2001) viable (MacDougall et al., 2001, LaJeunesse et al., 1998) visible (Fehon et al., 1997) visible \| recessive (LaJeunesse et al., 2001, LaJeunesse et al., 1998, McCartney et al., 2000)
Phenotype Manifest In
	adult cuticle & head (LaJeunesse et al., 1998) adult cuticle & head \| posterior (McCartney et al., 2000) adult cuticle & head \| ventral (McCartney et al., 2000) egg chamber \| posterior & follicle cell \| somatic clone \| rescuable maternal effect \| cell non-autonomous (MacDougall et al., 2001) elongation stage spermatid (Dorogova et al., 2008) eye (LaJeunesse et al., 2001, LaJeunesse et al., 1998, McCartney et al., 2000) eye \| anterior \| ventral (LaJeunesse et al., 2001) eye \| somatic clone (LaJeunesse et al., 1998) eye disc (McCartney et al., 2000) individualization stage spermatid (Dorogova et al., 2008) morphogenetic furrow (McCartney et al., 2000) morphogenetic furrow \| ectopic (McCartney et al., 2000) nucleus & elongation stage spermatid (Dorogova et al., 2008) ommatidium (LaJeunesse et al., 2001) oocyte nucleus (MacDougall et al., 2001) oocyte nucleus \| cell non-autonomous \| rescuable maternal effect \| somatic clone (MacDougall et al., 2001) photoreceptor cell \| ectopic (McCartney et al., 2000) posterior crossvein (LaJeunesse et al., 2001, McCartney et al., 2000) seminal vesicle (Dorogova et al., 2008) spermatocyte (Dorogova et al., 2008) spermatozoon (Dorogova et al., 2008) spindle & spermatocyte (Dorogova et al., 2008) vibrissae (LaJeunesse et al., 2001) vibrissae \| ectopic (McCartney et al., 2000) wing (LaJeunesse et al., 2001, LaJeunesse et al., 1998, McCartney et al., 2000) wing vein (LaJeunesse et al., 2001) wing vein L2 (LaJeunesse et al., 2001)
Detailed Description
	Statement Reference The seminal vesicles of mutant males are smaller than normal and are shrivelled. Very few sperm are found in the seminal vesicles and those that are found are immotile. The testes have fewer sperm heads in each bundle than normal and the bundles are disorganised. No abnormalities are found during mitosis of spermatogonia or spermatocyte growth in the mutant testes. Three types of abnormality are occasionally found during meiosis of spermatocytes. In approximately 5% of cases, spermatids containing two nuclei of equal size and two Nebenkerns are seen. Rarely, tripolar or four-polar spindles are seen in spermatocytes undergoing the second meiotic division. Spermatid nuclei in elongating cysts are grouped into two locations in some cysts and appear more scattered than normal in others. Some mutant spermatids at the late stage of spermatid elongation contain two paracrystalline bodies within the major mitochondrial derivative compared to one in control cysts. Some mutant spermatids have two axonemes present compared to only one in control cysts. Mutant cysts are loosely arranged compared to control cysts, which display cell-cell contact. Some cytoplasmic shedding is present mutant cysts. The sperm nuclei in mutant spermatid cysts undergoing individualization have variable morphology and appear scattered. The actin cones are dispersed. At the individualisation stage, mutant spermatids show variable phenotypes, including two paracrystalline body-filled mitochondrial derivatives with abnormal shape, or three paracrystalline body-filled mitochondrial derivatives together with one axoneme, or spermatids having one or two paracrystalline-filled mitochondrial derivatives but no axoneme compared to control spermatids. Mutant cysts show gross disorganization in the arrangement of the individualized spermatids. Some spermatids appear fused together. (Dorogova et al., 2008) Hemizygous males have broadened wings and show a low penetrance of disruptions of the posterior crossvein. The eyes are slightly rough and smaller than normal, and 10% have ectopic growths and vibrissae almost exclusively in the anterior ventral portion of the eye. Minor perturbation of ommatidial organisation is seen. Approximately 50% of the expected number of Mer³/Y males eclose. (LaJeunesse et al., 2001) Mosaic egg chambers in which all the follicle cells are homozygous, or in which a large homozygous clone covers all the posterior follicle cells show a strong Mer mutant phenotype indistinguishable from non-mosaic homozygous mutants. The oocyte nucleus fails to migrate. Mosaic egg chambers with homozygous clones in the anterior follicle cells or main body follicle cells do not have a Mer mutant phenotype. In mosaic egg chambers in which only a few posterior follicle cells are homozygous, the heterozygous cells rescue the Mer mutant phenotype in all the neighbouring follicle cells and also rescue the oocyte nuclear migration phenotype, indicating that Mer acts non-autonomously. (MacDougall et al., 2001) The ventral portion of the eye disc is significantly enlarged and the morphogenetic furrow appears to neither initiate or progress through the ectopic tissue in hemizygous animals. In less than 5% of cases, an ectopic morphogenetic furrow develops in the ventral portion of the disc with consequent neuronal differentiation. Hemizygous adults have a weakly roughened, reduced eye with expansion of ventral and posterior head cuticle and development of ectopic vibrissae (the bristles found ventral and anterior to the eye). There are only minor perturbations in interommatidial organisation and no obvious disruptions in ommatidial polarity. Hemizygous adults have broadened wing blades and may show disruption of the posterior crossvein. The increased wing size appears to be due to an increase in cell number rather than an increase in cell size and in fact cell size is slightly decreased compared to wild type. (McCartney et al., 2000) Homozygotes have broadened wings, and show a low and variably penetrant expression of weakly roughened eyes and development of abnormal head cuticle structures. Homozygous clones in the eye have a normal overall morphology, although occasionally a very weak roughened phenotype is seen. The clones are significantly larger than their wild-type sister clones. (LaJeunesse et al., 1998) Rescued by Mer^hs.PM. (Fehon et al., 1997)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhanced by
Statement Reference Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by E(Mer)IIb¹⁸² (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by E(Mer)IIb¹⁸⁷ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by E(Mer)IIb²⁰⁹ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by E(Mer)IIIa²³⁹ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by E(Mer)IIIa²⁷⁸ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by sbb⁹⁴ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by sbb¹⁵¹ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by sbb²¹⁶ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by sbb²⁵⁶ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by sbb²⁵⁹ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by sbb²⁷⁰ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, enhanceable by sbb³²⁴ (LaJeunesse et al., 2001) Mer³ has visible \| recessive phenotype, enhanceable by dpp^hr56/dpp[+] (McCartney et al., 2000) Mer³ has visible \| recessive phenotype, enhanceable by ex⁶⁹⁷ (McCartney et al., 2000) Mer³ has visible \| recessive phenotype, enhanceable by ex^e1 (McCartney et al., 2000) Mer³ has visible phenotype, enhanceable by +/In(2R)bw^VDe2LCy^R (LaJeunesse et al., 2001) Mer³ has visible phenotype, enhanceable by bs[+]/bs²⁴² (LaJeunesse et al., 2001) Mer³ has visible phenotype, enhanceable by bs[+]/bs²⁵³ (LaJeunesse et al., 2001) Mer³ has visible phenotype, enhanceable by bs³⁶⁴/bs[+] (LaJeunesse et al., 2001) Mer³ has visible phenotype, enhanceable by bs⁰³²⁶⁷/bs[+] (LaJeunesse et al., 2001) Mer³ has visible phenotype, enhanceable by Df(2L)C144/+ (LaJeunesse et al., 2001)
NOT Enhanced by
Statement Reference Mer³ has lethal \| male \| partially \| recessive phenotype, non-enhanceable by bs²⁴² (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, non-enhanceable by bs²⁵³ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, non-enhanceable by net¹⁰⁷ (LaJeunesse et al., 2001) Mer³ has lethal \| male \| partially \| recessive phenotype, non-enhanceable by net³⁸³ (LaJeunesse et al., 2001)
Suppressed by
Statement Reference Mer³ has visible phenotype, suppressible by bs[+]/bs²⁴² (LaJeunesse et al., 2001) Mer³ has visible phenotype, suppressible by bs[+]/bs²⁵³ (LaJeunesse et al., 2001) Mer³ has visible phenotype, suppressible by bs³⁶⁴/bs[+] (LaJeunesse et al., 2001) Mer³ has visible phenotype, suppressible by bs⁰³²⁶⁷/bs[+] (LaJeunesse et al., 2001) Mer³ has visible phenotype, suppressible by net[+]/net³⁸³ (LaJeunesse et al., 2001) Mer³ has visible phenotype, suppressible by net¹⁰⁷/net[+] (LaJeunesse et al., 2001)
Other
Statement Reference Df(2R)PC4/+, Mer³ has lethal phenotype (LaJeunesse et al., 2001)
Phenotype Manifest In
Enhanced by
Statement Reference Mer³ has adult cuticle & head phenotype, enhanceable by ex⁶⁹⁷ (McCartney et al., 2000) Mer³ has adult cuticle & head phenotype, enhanceable by ex^e1 (McCartney et al., 2000) Mer³ has adult head & cuticle phenotype, enhanceable by dpp^hr56/dpp[+] (McCartney et al., 2000) Mer³ has eye phenotype, enhanceable by +/In(2R)bw^VDe2LCy^R (LaJeunesse et al., 2001) Mer³ has eye phenotype, enhanceable by Df(2L)C144/+ (LaJeunesse et al., 2001) Mer³ has eye phenotype, enhanceable by dpp^hr56/dpp[+] (McCartney et al., 2000) Mer³ has eye phenotype, enhanceable by ex⁶⁹⁷ (McCartney et al., 2000) Mer³ has interommatidial bristle \| ectopic phenotype, enhanceable by ex⁶⁹⁷ (McCartney et al., 2000) Mer³ has interommatidial bristle phenotype, enhanceable by dpp^hr56/dpp[+] (McCartney et al., 2000) Mer³ has interommatidial bristle phenotype, enhanceable by ex⁶⁹⁷ (McCartney et al., 2000) Mer³ has ommatidium phenotype, enhanceable by +/In(2R)bw^VDe2LCy^R (LaJeunesse et al., 2001) Mer³ has ommatidium phenotype, enhanceable by Df(2L)C144/+ (LaJeunesse et al., 2001) Mer³ has ommatidium phenotype, enhanceable by dpp^hr56/dpp[+] (McCartney et al., 2000) Mer³ has posterior crossvein phenotype, enhanceable by bs²⁴² (LaJeunesse et al., 2001) Mer³ has vibrissae \| ectopic phenotype, enhanceable by ex⁶⁹⁷ (McCartney et al., 2000) Mer³ has vibrissae phenotype, enhanceable by +/In(2R)bw^VDe2LCy^R (LaJeunesse et al., 2001) Mer³ has vibrissae phenotype, enhanceable by Df(2L)C144/+ (LaJeunesse et al., 2001)
NOT Enhanced by
Statement Reference Mer³ has head phenotype, non-enhanceable by bs²⁴² (LaJeunesse et al., 2001) Mer³ has head phenotype, non-enhanceable by bs²⁵³ (LaJeunesse et al., 2001) Mer³ has head phenotype, non-enhanceable by net¹⁰⁷ (LaJeunesse et al., 2001) Mer³ has head phenotype, non-enhanceable by net³⁸³ (LaJeunesse et al., 2001) Mer³ has leg phenotype, non-enhanceable by bs²⁴² (LaJeunesse et al., 2001) Mer³ has leg phenotype, non-enhanceable by bs²⁵³ (LaJeunesse et al., 2001) Mer³ has leg phenotype, non-enhanceable by net¹⁰⁷ (LaJeunesse et al., 2001) Mer³ has leg phenotype, non-enhanceable by net³⁸³ (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by +/Df(2L)Mdh (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by Df(2L)cact-255rv64/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by Df(2L)H20/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by Df(2L)net-PMF/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by Df(2R)cn88b/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by Df(2R)en30/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by Df(3R)Antp17/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by Df(3R)awd-KRB/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by Df(3R)Tl-P/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by sbb⁰⁴⁴⁴⁰ (LaJeunesse et al., 2001) Mer³ has phenotype, non-enhanceable by sbb^k00702 (LaJeunesse et al., 2001)
Suppressed by
Statement Reference Mer³ has wing phenotype, suppressible \| partially by bs[+]/bs²⁴² (LaJeunesse et al., 2001) Mer³ has wing phenotype, suppressible \| partially by bs[+]/bs²⁵³ (LaJeunesse et al., 2001) Mer³ has wing phenotype, suppressible \| partially by bs³⁶⁴/bs[+] (LaJeunesse et al., 2001) Mer³ has wing phenotype, suppressible \| partially by bs⁰³²⁶⁷/bs[+] (LaJeunesse et al., 2001) Mer³ has wing phenotype, suppressible \| partially by net[+]/net³⁸³ (LaJeunesse et al., 2001) Mer³ has wing phenotype, suppressible \| partially by net¹⁰⁷/net[+] (LaJeunesse et al., 2001) Mer³ has wing vein phenotype, suppressible by net³⁸³ (LaJeunesse et al., 2001)
NOT suppressed by
Statement Reference Mer³ has head phenotype, non-suppressible by bs²⁴² (LaJeunesse et al., 2001) Mer³ has head phenotype, non-suppressible by bs²⁵³ (LaJeunesse et al., 2001) Mer³ has head phenotype, non-suppressible by net¹⁰⁷ (LaJeunesse et al., 2001) Mer³ has head phenotype, non-suppressible by net³⁸³ (LaJeunesse et al., 2001) Mer³ has leg phenotype, non-suppressible by bs²⁴² (LaJeunesse et al., 2001) Mer³ has leg phenotype, non-suppressible by bs²⁵³ (LaJeunesse et al., 2001) Mer³ has leg phenotype, non-suppressible by net¹⁰⁷ (LaJeunesse et al., 2001) Mer³ has leg phenotype, non-suppressible by net³⁸³ (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by +/Df(2L)Mdh (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by Df(2L)cact-255rv64/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by Df(2L)H20/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by Df(2L)net-PMF/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by Df(2R)cn88b/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by Df(2R)en30/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by Df(3R)Antp17/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by Df(3R)awd-KRB/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by Df(3R)Tl-P/+ (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by sbb⁰⁴⁴⁴⁰ (LaJeunesse et al., 2001) Mer³ has phenotype, non-suppressible by sbb^k00702 (LaJeunesse et al., 2001)
Additional Comments
Genetic Interactions
Statement Reference Mutant phenotype is dominantly modified by Df(2L)al, Df(2L)ast2, Df(2L)JS32, Df(2L)cl-h3, Df(2L)30A-C, Df(2L)J39, Df(2R)M41A, Df(2R)nap1, Df(2R)pk78s, Df(2R)44CE, Df(2R)en-A, Df(2R)or-BR6, Df(3L)HR119, Df(3L)ZN47, Df(3L)brm11, Df(3L)VW3, Df(3R)T-32, Df(3R)ry615, Df(3R)P14, Df(3R)Cha7 or Df(3R)e-N19. Df(2L)C144 or In(2R)bw^VDe2LCy^R dominantly enhances the Mer³ phenotypes, including reduction in size of the eye and formation of aberrant cuticle, bristles and outgrowths. The increase in the size of the wing blade seen in Mer³/Y flies is partially suppressed by bs²⁵³/+, bs²⁴²/+, bs³⁶⁴/+, bs⁰³²⁶⁷/+, net¹⁰⁷/+ or net³⁸³/+. (LaJeunesse et al., 2001) dpp^hr56 dominantly enhances the phenotype of Mer³ hemizygotes; the eye is smaller and more roughened and there is increased expansion of peripheral head cuticle. Numerous interommatidial bristle duplications are seen. (McCartney et al., 2000)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Rescued by	Mer³ is rescued by Mer^hs.PM (Fehon et al., 1997)
Comments
Stocks ( 1 )
Bloomington	9103 w^* Mer³ P{neoFRT}19A/FM6
Notes on Origin
Discoverer

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 1 )
Reported As
Symbol Synonym	Mer³ (Hawley and Gilliland, 2006, Dorogova et al., 2008, Popodi et al., 2010-)
Name Synonym
Secondary FlyBase IDs

References ( 9 )
Research paper	Dorogova et al., 2008, BMC Cell Biol. 9: 1 The role of Drosophila Merlin in spermatogenesis. [FBrf0205210] LaJeunesse et al., 2001, Genetics 158(2): 667--679 A Systematic screen for dominant second-site modifiers of Merlin/NF2 Phenotypes reveals an interaction with blistered/DSRF and scribbler. [FBrf0136951] MacDougall et al., 2001, Development 128(5): 665--673 Merlin, the Drosophila homologue of neurofibromatosis-2, is specifically required in posterior follicle cells for axis formation in the oocyte. [FBrf0134575] McCartney et al., 2000, Development 127(6): 1315--1324 The Neurofibromatosis-2 homologue, Merlin, and the tumor suppressor expanded function together in Drosophila to regulate cell proliferation and differentiation. [FBrf0125175] LaJeunesse et al., 1998, J. Cell Biol. 141(7): 1589--1599 Structural analysis of Drosophila Merlin reveals functional domains important for growth control and subcellular localization. [FBrf0102961] Fehon et al., 1997, Genetics 146(1): 245--252 Isolation of mutations in the Drosophila homologues of the human Neurofibromatosis 2 and yeast CDC42 genes using a simple and efficient reverse-genetic method. [FBrf0093443]
Review	Hawley and Gilliland, 2006, Genetics 174(1): 5--15 Sometimes the result is not the answer: The truths and the lies that come from using the complementation test. [FBrf0194489]
Personal communication to FlyBase	Popodi et al., 2010-, Small X duplications for the stock center collection. Small X duplications for the stock center collection. [FBrf0210621]
Abstract	McCartney and Fehon, 1998, A. Dros. Res. Conf. 39: 298B Loss of Function of the Neurofibromatosis 2 Tumor Suppressor Merlin Leads to Overproliferation in Drosophila. [FBrf0101335]

Allele Dmel\Mer3

Allele Dmel\Mer³