Allele Dmel\ru¹

General Information
Symbol	Dmel\ru¹	Species	D. melanogaster
Name		FlyBase ID	FBal0014832
Feature type	allele	Created / Updated	2006-08-22/2006-08-22
Associated gene	Dmel\ru

Allele class	hypomorph
Mutagen	spontaneous
Nature of the Allele
Allele class	hypomorph (Wasserman et al., 2000)
Mutagen	spontaneous
Mapped Features and Mutations
Type Symbol & 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
Assay mode
Carried on aberration	C(3L)RM-VG1 DcxF-Sse (FlyBase Curators, 2006-) DcxF (FlyBase Curators, 2006-) TM3-mwh²-ru¹ (FlyBase Curators, 2006-, Roote and Ashburner, 2005.6)
Cytology
Phenotypic Data
Phenotypic Class
	cell polarity defective (Brown and Freeman, 2003)
Phenotype Manifest In
	ommatidium (Brown and Freeman, 2003) photoreceptor (Brown and Freeman, 2003) embryonic/larval ganglionic branch (Gallio et al., 2004) eye (Wasserman et al., 2000) cone cell (Wasserman et al., 2000) pigment cell (Wasserman et al., 2000)
Detailed Description
	Statement Reference may overlap wild type Eyes small and rough, have irregular facets and hairs and have black specks from erupted facets. Expression variable; sometimes overlaps wild type. SEM study by Stumm-Tegethoff and Dick (1974). The regular array of facets in the eyes of homozygotes produces a rough eye phenotype. There is variable loss of photoreceptors in these mutant eyes, however few defects if any are seen in the recruitment of photoreceptors. Cone cells are dramatically under-recruited, and some pigment cells are seen to be missing. (Wasserman et al., 2000) Heterozygous adults exhibit misrotated ommatidia and minor photoreceptor recruitment defects. (Brown and Freeman, 2003) In stage 16 ru¹ mutant embryos, a significant number of ganglionic branch trachea fail to turn posteriorly and dorsally at the ventral midline and instead cross the midline or remain lingering on it. (Gallio et al., 2004)
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhanced by
Statement Reference ru¹ has visible phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000)
Enhancer of
Statement Reference ru¹ is an enhancer of visible phenotype of rho^7M43 (Wasserman et al., 2000)
NOT Enhancer of
Statement Reference ru¹ is a non-enhancer of cell polarity defective phenotype of nmo^P1 (Brown and Freeman, 2003)
NOT Suppressor of
Statement Reference ru¹ is a non-suppressor of cell polarity defective phenotype of nmo^P1 (Brown and Freeman, 2003)
Other
Statement Reference rho^7M43, ru¹ has cell lethal \| somatic clone phenotype (Wasserman et al., 2000, Wasserman et al., 2000) rho^7M43, ru¹ has cell death defective \| somatic clone phenotype (Wasserman et al., 2000, Wasserman et al., 2000) rho^7M43, ru¹ has mitotic cell cycle defective \| somatic clone phenotype (Yang and Baker, 2003, Yang and Baker, 2003) rho^7M43, ru¹ has mitotic cell cycle defective \| somatic clone \| cell non-autonomous phenotype (Yang and Baker, 2003, Yang and Baker, 2003) rho^7M43, ru¹ has cell death increase \| embryonic stage phenotype (Parker, 2006) rho^7M43, ru¹, vn^L6 has visible phenotype (Campbell, 2002, Campbell, 2002, Campbell, 2002) Df(3L)ru-K1, ru¹ has visible phenotype (Krivshenko, 1958)
Phenotype Manifest In
Enhanced by
Statement Reference ru¹ has cone cell \| somatic clone phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) ru¹ has eye \| somatic clone phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) ru¹ has photoreceptor cell R1 \| somatic clone phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) ru¹ has photoreceptor cell R2 \| somatic clone phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) ru¹ has photoreceptor cell R3 \| somatic clone phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) ru¹ has photoreceptor cell R4 \| somatic clone phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) ru¹ has photoreceptor cell R5 \| somatic clone phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) ru¹ has photoreceptor cell R6 \| somatic clone phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) ru¹ has photoreceptor cell R7 \| somatic clone phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) ru¹ has photoreceptor cell R8 \| somatic clone \| cell non-autonomous phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) ru¹ has photoreceptor cell \| somatic clone phenotype, enhanceable by rho^7M43 (Wasserman et al., 2000) rho^7M43, ru¹ has photoreceptor cell \| somatic clone phenotype, enhanceable by sens^E2 (Frankfort and Mardon, 2004)
Enhancer of
Statement Reference ru¹ is an enhancer of cone cell \| somatic clone phenotype of rho^7M43 (Wasserman et al., 2000) ru¹ is an enhancer of eye \| somatic clone phenotype of rho^7M43 (Wasserman et al., 2000) ru¹ is an enhancer of photoreceptor cell R1 \| somatic clone phenotype of rho^7M43 (Wasserman et al., 2000) ru¹ is an enhancer of photoreceptor cell R2 \| somatic clone phenotype of rho^7M43 (Wasserman et al., 2000) ru¹ is an enhancer of photoreceptor cell R3 \| somatic clone phenotype of rho^7M43 (Wasserman et al., 2000) ru¹ is an enhancer of photoreceptor cell R4 \| somatic clone phenotype of rho^7M43 (Wasserman et al., 2000) ru¹ is an enhancer of photoreceptor cell R5 \| somatic clone phenotype of rho^7M43 (Wasserman et al., 2000) ru¹ is an enhancer of photoreceptor cell R6 \| somatic clone phenotype of rho^7M43 (Wasserman et al., 2000) ru¹ is an enhancer of photoreceptor cell R7 \| somatic clone phenotype of rho^7M43 (Wasserman et al., 2000) ru¹ is an enhancer of photoreceptor cell R8 \| somatic clone \| cell non-autonomous phenotype of rho^7M43 (Wasserman et al., 2000) ru¹ is an enhancer of photoreceptor cell \| somatic clone phenotype of rho^7M43 (Wasserman et al., 2000)
NOT Enhancer of
Statement Reference ru¹ is a non-enhancer of ommatidium phenotype of nmo^P1 (Brown and Freeman, 2003)
NOT Suppressor of
Statement Reference ru¹ is a non-suppressor of ommatidium phenotype of nmo^P1 (Brown and Freeman, 2003)
Other
Statement Reference rho^7M43, ru¹ has photoreceptor cell R2 \| somatic clone phenotype (Yang and Baker, 2003, Yang and Baker, 2003, Frankfort and Mardon, 2004, Frankfort and Mardon, 2004) rho^7M43, ru¹ has photoreceptor cell R3 \| somatic clone phenotype (Yang and Baker, 2003, Yang and Baker, 2003, Frankfort and Mardon, 2004, Frankfort and Mardon, 2004) rho^7M43, ru¹ has photoreceptor cell R4 \| somatic clone phenotype (Yang and Baker, 2003, Yang and Baker, 2003, Frankfort and Mardon, 2004, Frankfort and Mardon, 2004) rho^7M43, ru¹ has photoreceptor cell R5 \| somatic clone phenotype (Yang and Baker, 2003, Yang and Baker, 2003, Frankfort and Mardon, 2004, Frankfort and Mardon, 2004) rho^7M43, ru¹ has photoreceptor cell R1 \| somatic clone phenotype (Frankfort and Mardon, 2004, Frankfort and Mardon, 2004) rho^7M43, ru¹ has photoreceptor cell R6 \| somatic clone phenotype (Frankfort and Mardon, 2004, Frankfort and Mardon, 2004) rho^7M43, ru¹ has photoreceptor cell R7 \| somatic clone phenotype (Frankfort and Mardon, 2004, Frankfort and Mardon, 2004) rho^7M43, ru¹ has ommatidial cluster \| somatic clone phenotype (Brown et al., 2006) rho^7M43, ru¹, vn^L6 has tarsal segment phenotype (Campbell, 2002, Campbell, 2002, Campbell, 2002) Df(3L)ru-K1, ru¹ has eye phenotype (Krivshenko, 1958) Df(3L)ru-K1, ru¹ has ommatidium phenotype (Krivshenko, 1958) Gug¹⁴⁹⁶⁷, argos^Δ7, rho^7M43, ru¹ has eye photoreceptor cell \| somatic clone phenotype (Charroux et al., 2006, Charroux et al., 2006, Charroux et al., 2006, Charroux et al., 2006)
Additional Comments
Genetic Interactions
Statement Reference ru¹/Df(3L)ru-K1 flies have slightly diminished eyes and the facets are rough. (Krivshenko, 1958) Clones of doubly mutant for rho^7M43 and ru¹ do not survive into adult eyes. When mosaic ommatidia are studied, no mutant R8 photoreceptor cells are seen, while mutant photoreceptor cells R1-7 are seen in between a third and a half of mosaic ommatidia. When imaginal discs are examined, an absence of non-R8 photoreceptor cells are seen, and also a complete loss of cone cells. In these clones an increased level of apoptosis is also seen. Apoptotic cells are seen in two main zones, one just ahead of the advancing morphogenetic furrow, and one towards the posterior of the clone. (Wasserman et al., 2000) Large ru¹ rho^7M43 vn^L6 triple mutant clones in the leg can result in truncations of the tarsus region (for example only three tarsal segments may be present). Mosaic legs which have wild-type tissue at the distal tip show rescue of tarsal development. (Campbell, 2002) In rho^7M43; ru¹ double mutant somatic clones in the 3rd instar eye disc, photoreceptors R2-R5 fail to undergo or differentiation. Within rho^7M43; ru¹ double mutant somatic clones there is a failure of G1 arrest in the furrow: all cells except R8s re-enter the cell cycle. Most of these cells fail to progress past G2: most remain in G2 arrest posterior to column 3, but both cells in early mitosis and postmitotic cells can be seen near boundaries with wild-type cells. (Yang and Baker, 2003) In rho^7M43; ru¹ double mutant clones in the eye disc, only R8 photoreceptor cells differentiate In rho^7M43; ru¹; sens^E2 triple mutant clones, no photoreceptors differentiate except for a few photoreceptors near the clonal boundary, presumably rescued non-autonomously by neighboring wild-type cell. No rescue of photoreceptor development is seen when these triple mutant clones are made in a ro^x63 homozygous background. (Frankfort and Mardon, 2004) 69% of cuticles from rho^7M43; ru¹ double homozygous embryos have at least one denticle belt fusion, compared with 30% in rho^unspecified homozygotes. Other aspects of the cuticle phenotype in these double mutants are no stronger than those in rho^unspecified embryos. (Note, while the authors do not name an rho allele for this analysis, they do claim to have used a null allele.) Denticle belt fusions in the cuticles of rho^7M43; ru¹ double homozygous embryos are suppressed by Egfr::tor^{t4021E.hs.sev} or Ras85D^V12.Scer\UAS with Scer\GAL4^prd.RG1, and enhanced by Egfr^DN.Scer\UAS (P{UAS-Egfr.DN}29-77-1), Ras85D^N17.Scer\UAS or phl^{K497M.Scer\UAS} with Scer\GAL4^prd.RG1. The penetrance of denticle belt fusions in the cuticles of rho^7M43; ru¹ double homozygous embryos is reduced from just under 60% to less than 20% by Df(3L)H99/Df(3L)H99. (Urban et al., 2004) ru[1], rho[7M43] embryos show an increased level of apoptosis. (Parker, 2006) The extra photoreceptor cell phenotype of argos[Δ7] clones is enhanced in argos[Δ7] Gug[14967] clones, with a greater percentage of ommatidia showing supernumerary photoreceptor cells. In contrast, ru[1] rho[7M43] Gug[14967] argos[Δ7] mutant clones lack photoreceptor cells. (Charroux et al., 2006) ru[1] rho[7M43] Minute clones in the eye disc fail to form the arcs and rosettes of cells in the morphogenetic furrow that are seen in wild-type eye discs. Small clusters of cells can very occasionally be observed in the furrow but these appear at much lower levels than in wild type. (Brown et al., 2006)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Rescued by	ru¹ is rescued by ru^hs.PW (Wasserman et al., 2000)
Comments
Stocks ( 248 )
Bloomington	575 ru¹ 3256 ru¹ klar¹ 1116 C(2)EN, bw¹; ru¹ 581 ru¹ lxd¹ by¹ 6882 ru¹ Aprt⁵ h¹ 4499 Df(3L)Scf-R11, ru¹/TM3, Sb¹ 600 Df(3L)Aprt-1, ru¹ h¹/TM3, Ser¹ 1257 ru¹ h¹ Dew¹/TM3, Ser¹ 93 ru¹ h¹ st¹ ry⁵⁰⁶ e¹
Kyoto	106944 ru¹ klar¹ 107762 Df(3L)Scf-R11, ru¹/TM3, Sb¹ 105729 ru¹ h¹ st¹ ry⁵⁰⁶ e¹
	More stocks available...
Notes on Origin
Discoverer	Sturtevant, 14th Feb. 1919.

Synonyms & Secondary IDs ( 3 )
Reported As
Symbol Synonym	rho-3¹ (Frankfort and Mardon, 2004) ru (Krivshenko, 1958) ru¹ (Parker, 2006, Charroux et al., 2006, Brown et al., 2006)
Name Synonym
Secondary FlyBase IDs

References ( 16 )
Research paper	Brown et al., 2006, Dev. Biol. 300(2): 710--721 Epithelial cell adhesion in the developing Drosophila retina is regulated by Atonal and the EGF receptor pathway. [FBrf0192868] Charroux et al., 2006, Dev. Biol. 291(2): 278--290 Atrophin contributes to the negative regulation of epidermal growth factor receptor signaling in Drosophila. [FBrf0190100] Parker, 2006, Curr. Biol. 16(20): 2058--2065 Control of compartment size by an EGF ligand from neighboring cells. [FBrf0193620] Frankfort and Mardon, 2004, Development 131(3): 563--570 Senseless represses nuclear transduction of Egfr pathway activation. [FBrf0167533] Gallio et al., 2004, Development 131(15): 3605--3614 Rhomboid 3 orchestrates Slit-independent repulsion of tracheal branches at the CNS midline. [FBrf0179216] Urban et al., 2004, Development 131(8): 1835--1845 EGF receptor signalling protects smooth-cuticle cells from apoptosis during Drosophila ventral epidermis development. [FBrf0174566] Brown and Freeman, 2003, Development 130(22): 5401--5412 Egfr signalling defines a protective function for ommatidial orientation in the Drosophila eye. [FBrf0167498] Yang and Baker, 2003, Dev. Cell 4(3): 359--369 Cell cycle withdrawal, progression, and cell survival regulation by EGFR and its effectors in the differentiating Drosophila eye. [FBrf0158858] Campbell, 2002, Nature 418(6899): 781--785 Distalization of the Drosophila leg by graded EGF-receptor activity. [FBrf0151696] Wasserman et al., 2000, Genes Dev. 14(13): 1651--1663 A family of rhomboid-like genes: Drosophila rhomboid-1 and roughoid/rhomboid-3 cooperate to activate EGF receptor signaling. [FBrf0128677] Krivshenko, 1958, D. I. S. 32: 80--81 [New mutants report.] [FBrf0063600] Bridges and Morgan, 1923, Publs Carnegie Instn 327: 1--251 The third-chromosome group of mutant characters of Drosophila melanogaster. [FBrf0001261] Strong, 1920, Biol. Bull., Wood's Hole 38: 33--37 Roughoid, a mutant located to the left of sepia in the third chromosome of Drosophila melanogaster. [FBrf0001011]
Personal communication to FlyBase	Roote and Ashburner, 2005.6, mwh<up>2</up>. mwh<up>2</up>. [FBrf0188644]
FlyBase analysis	FlyBase Curators, 2006-, Balancer summary information. Balancer summary information. [FBrf0189689]
Book	Morgan et al., 1925, Biblphia Genet. 2: 262pp The genetics of Drosophila melanogaster. [FBrf0001379]

Allele Dmel\ru1

Allele Dmel\ru¹