Allele Dmel\fu⁵²

General Information
Symbol	Dmel\fu⁵²	Species	D. melanogaster
Name		FlyBase ID	FBal0004885
Feature type	allele	Associated gene	Dmel\fu
Also Known As	fu^JB3

Map ( GBrowse )
Allele class	hypomorphic allele - genetic evidence
Mutagen	diepoxybutane
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	hypomorphic allele - genetic evidence (Glise et al., 2002)
Mutagen	diepoxybutane (Busson et al., 1988, Preat et al., 1993, Therond et al., 1996)
Mutations Mapped to the Genome

Type Location Additional Notes References deletion X:18,551,971..18,552,010 comment=40bp deletion evidence=experimental (Therond et al., 1996)
Associated Sequence Data
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name

UniProtKB/Swiss-Prot
UniProtKB/TrEMBL

Progenitor genotype
Nature of the lesion	Statement Reference Defined by the absence of residues 29 to 37 in the N-terminal catalytic domain. (Daoud and Blanchet-Tournier, 2005) 40bp deletion of genomic sequences 1002-1041, deletion of the 3' part of the first exon and 5' part of the first intron. (Therond et al., 1996) Mutation in the kinase domain. Class I fu allele, affects the catalytic domain but does not change the open reading frame. (Preat et al., 1993) Small deletion within putative kinase domain. (Preat et al., 1990)
Cytology
Phenotypic Data
Phenotypic Class
	female sterile tumorigenic (Narbonne-Reveau et al., 2006) tumorigenic (with Df(1)fu-Z4) (Narbonne-Reveau et al., 2006) tumorigenic (with fu⁵⁰) (Narbonne-Reveau et al., 2006) viable \| reduced (Busson et al., 1988) visible (Glise et al., 2002, Busson et al., 1988) visible \| recessive (Daoud and Blanchet-Tournier, 2005)
Phenotype Manifest In
	cystoblast \| supernumerary (Narbonne-Reveau et al., 2006) cystoblast \| supernumerary (with Df(1)fu-Z4) (Narbonne-Reveau et al., 2006) cystoblast \| supernumerary (with fu⁵⁰) (Narbonne-Reveau et al., 2006) discal cell (Daoud and Blanchet-Tournier, 2005) egg chamber (Besse et al., 2002, Narbonne-Reveau et al., 2006) egg chamber (with Df(1)fu-Z4) (Narbonne-Reveau et al., 2006) egg chamber (with fu⁵⁰) (Narbonne-Reveau et al., 2006) interfollicle cell (Besse et al., 2002) interfollicle cell \| somatic clone (Besse et al., 2002) nurse cell (Narbonne-Reveau et al., 2006) oocyte (Narbonne-Reveau et al., 2006) ovariole (Besse et al., 2002) polar follicle cell (Besse et al., 2002, Besse and Pret, 2003) wing wing basal cell 2 (Daoud and Blanchet-Tournier, 2005) wing vein L3 (Busson et al., 1988, Daoud and Blanchet-Tournier, 2005) wing vein L3 \| distal (Glise et al., 2002) wing vein L3 \| proximal (Glise et al., 2002) wing vein L4 (Busson et al., 1988, Daoud and Blanchet-Tournier, 2005)
Detailed Description
	Statement Reference 4 day old fu[52] homozygous females, only 11% of egg chambers are tumorous, increasing to 35% at 10 days. This phenotype is stronger in fu[50]/fu[52] females, in which 83% of egg chambers are tumorous ay 4 days, decreasing to 71% at 10 days. This is stronger than the phenotype in fu[52]/Df(1)fu-Z4 females, where around 50% of egg chambers are tumorous at 4 or 10 days. Tumorous follicles in these ovarioles consist of tens to hundreds of germ-line cells with small non-polyploid nuclei enveloped by a regular follicular epithelium. Marker expression studies and analysis of spectrosome morphology in tumorous cysts from fu[52]/Df(1)fu-Z4 animals suggest that these tumorous germ-line cells remain very immature and do not progress beyond the early cystoblast stage and may include germline stem cells (GSCs). However, the size of germline stem cell niches appears to be unnaffected - suggesting that these phenotypes are due to inceases in cystoblast-like cells outside of the niche. This increase is not due to increased cell division of germline stem cells in the niche - as assayed by scoring the percentage of GSCs in the niche undergoing mitosis in fu[52]/fu[52] fu[52]/fu[50] and fu[52]/Df(1)fu-Z4 adult females. fu[52] homozygous ovaries also contain egg chambers with increased numbers of germ cells that, based on marker expression and fusome morphology, have developed past the cystolast stage but are still more immature than normally found in female germline cysts outside of the germarium. There is some evidence of oocytes and nurse cell determination, but nurse cell and oocyte differentation appear to be blocked. (Narbonne-Reveau et al., 2006) fu⁵²/+ heterozygotes have a partial fusion of wing veins L3 and L4. (Daoud and Blanchet-Tournier, 2005) Homozygous females show a delay in the polar cell differentiation program. Restriction in the final number of polar cells is achieved by stage 5 in these females. (Besse and Pret, 2003) Mutant ovaries contain some egg chambers with more or less than the normal number of 16 germline cells. When an egg chamber contains less than 15 nurse cells, the complementary nurse cells are sometimes found in an adjacent egg chamber. In one case, an egg chamber containing two oocytes has been observed, indicating a multicyst egg chamber. In some cases the follicular epithelia of two adjacent egg chambers are apposed with no interfollicular stalk between them. The same ovariole can contain both normal and abnormal egg chambers. Degenerating egg chambers are seen. More dividing cells are seen in germarial regions 2 and 3 than in wild type. Mutant germaria show impaired encapsulation of the germline cysts by prefollicular cells, which results in some multicyst chambers. Interfollicular stalk cells do appear to be specified in the mutant germaria, but they are unable to fulfill their role in budding off of individual egg chambers. When stalk-like structures are seen between egg chambers they have an abnormal morphology, consisting of aggregates of round cells arranged in a ball shape (instead of the wild-type linear arrangement of 5-7 oval shaped cells). There is a delay in polar follicle cell specification. Clones induced in the somatic stem cells of the ovarioles can result in long, disorganised stalks within the ovariole, from which egg chambers show an off centre attachment. In most cases, these stalks are composed of fu⁺ cells, while the mutant cells are found as part of the follicular epithelium of adjacent chambers. (Besse et al., 2002) Mutants show fusion of wing veins L3 and L4 both proximally and distally, with intervein tissue remaining between the two veins in the medial part of the wing. (Glise et al., 2002) Weak wing phenotype (LV3 and LV4 fused proximally), reduced viability, strong fecundity phenotype. (Busson et al., 1988) Weak wing phenotype. Weak viability phenotype.
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Phenotype Manifest In
Enhanced by
Statement Reference fu⁵² has discal cell phenotype, enhanceable by Scer\GAL4^ptc-559.1/Hsap\STK36^{Scer\UAS.Δ77-98} (Daoud and Blanchet-Tournier, 2005) fu⁵² has wing basal cell 2 phenotype, enhanceable by Scer\GAL4^ptc-559.1/Hsap\STK36^{Scer\UAS.Δ77-98} (Daoud and Blanchet-Tournier, 2005) fu⁵² has wing vein L3 phenotype, enhanceable by Scer\GAL4^ptc-559.1/Hsap\STK36^{Scer\UAS.Δ77-98} (Daoud and Blanchet-Tournier, 2005) fu⁵² has wing vein L4 phenotype, enhanceable by Scer\GAL4^ptc-559.1/Hsap\STK36^{Scer\UAS.Δ77-98} (Daoud and Blanchet-Tournier, 2005)
Suppressed by
Statement Reference fu⁵² has ovariole phenotype, suppressible \| partially by ci^{Ce-2.Scer\UAS.T:Ivir\HA1}/Scer\GAL4^Act5C.PP (Besse et al., 2002) fu⁵² has ovariole phenotype, suppressible \| partially by cos[+]/cos¹ (Besse et al., 2002) fu⁵² has ovariole phenotype, suppressible \| partially by Su(fu)^LP/Su(fu)^LP (Besse et al., 2002) fu⁵² has phenotype, suppressible \| partially by Su(fu)^LP (Preat et al., 1993)
Additional Comments
Genetic Interactions
Statement Reference Hsap\fu^{Scer\UAS.Δ77-98}; Scer\GAL4^ptc-559.1 enhances the partial fusion of wing veins L3 and L4 seen in fu⁵² flies. (Daoud and Blanchet-Tournier, 2005) When in combination with Su(fu)^LP heterozygotes individuals display a partially suppressed fu phenotype. When in combination with cos³ individuals display no cos phenotype. (Preat et al., 1993)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Comments
Stocks ( 1 )
Bloomington	7205 fu⁵²/FM6
Notes on Origin
Discoverer

Comments
	Class I mutation based on interaction with Su(fu)^LP, suppression of embryonic and adult phenotype. (Therond et al., 1996)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 2 )
Reported As
Symbol Synonym	fu⁵² fu^JB3 (Daoud and Blanchet-Tournier, 2005, Besse and Pret, 2003, Besse et al., 2002, Glise et al., 2002, Therond et al., 1996, Preat et al., 1993, Preat et al., 1990, Busson et al., 1988, Dussillol-Godar et al., 2006, Xia et al., 2012, Narbonne-Reveau et al., 2006)
Name Synonym
Secondary FlyBase IDs

References ( 11 )
Research paper	Xia et al., 2012, Curr. Biol. 22(6): 515--521 The Niche-Dependent Feedback Loop Generates a BMP Activity Gradient to Determine the Germline Stem Cell Fate. [FBrf0217877] Dussillol-Godar et al., 2006, Dev. Biol. 291(1): 53--66 Modulation of the Suppressor of fused protein regulates the Hedgehog signaling pathway in Drosophila embryo and imaginal discs. [FBrf0190115] Narbonne-Reveau et al., 2006, Mech. Dev. 123(3): 197--209 fused regulates germline cyst mitosis and differentiation during Drosophila oogenesis. [FBrf0191131] Daoud and Blanchet-Tournier, 2005, Dev. Genes Evol. 215(5): 230--237 Expression of the human FUSED protein in Drosophila. [FBrf0187435] Besse and Pret, 2003, Development 130(5): 1017--1027 Apoptosis-mediated cell death within the ovarian polar cell lineage of Drosophila melanogaster. [FBrf0155695] Besse et al., 2002, Development 129(17): 4111--4124 Fused-dependent Hedgehog signal transduction is required for somatic cell differentiation during Drosophila egg chamber formation. [FBrf0151279] Glise et al., 2002, Dev. Biol. 248(1): 93--106 Notch and wingless modulate the response of cells to hedgehog signalling in the Drosophila wing. [FBrf0151253] Therond et al., 1996, Genetics 142(4): 1181--1198 Functional domains of fused, a serine-threonine kinase required for signaling in Drosophila. [FBrf0087739] Preat et al., 1993, Genetics 135(4): 1047--1062 Segmental polarity in Drosophila melanogaster: genetic dissection of fused in a suppressor of fused background reveals interaction with costal-2. [FBrf0065539] Preat et al., 1990, Nature 347(4): 87--89 A putative serine/threonine protein kinase encoded by the segment-polarity fused gene of Drosophila. [FBrf0052668] Busson et al., 1988, Rouxs Arch. Dev. Biol. 197(4): 221--230 Genetic analysis of viable and lethal fused mutants of Drosophila melanogaster. [FBrf0048987]

Allele Dmel\fu52

Allele Dmel\fu⁵²