# r: rudimentary location: 1-54.5 [Revised to 55.6 by Colaianne and Bell (1972, DIS 48: 20-21) and to 55.3-55.4 by N/rby (1970, DIS 45: 41) based on large scale measurements of r-f recombination]. r9: rudimentary-9 From Edith M. Wallace, unpublished. phenotype: Homozygotes and hemizygotes are pyrimidine auxo- trophs (N|rby, 1969, Hereditas 66: 205-14). A complex locus encoding a 220 kd polypeptide containing the first three enzyme activities in the pyrimidine synthetic pathway: glutamine-dependent carbamyl phosphate synthetase [CPS (EC.2.7.2.5)] (Jarry and Falk, 1974, Mol. Gen. Genet. 135: 113-22), aspartate transcarbamylase [ATC (EC.2.1.3.2)] (N/rby, 1969), and dihydroorotase [DHO (EC.3.5.2.3)] (Rawls and Fristrom, 1975, Nature 255: 738-40). Probably exists as a homomultimer. These three activities cosediment and copurify (Brothers, Tsubota, Germeraad, and Fristrom, 1978, Biochem. Genet. 16: 321-32); also the developmental profiles of the three activities are the same, maximal in the egg, dropping until the time of hatching, increasing again during the first larval instar, and then leveling off at a low level (Mehl and Jarry, 1978, Dev. Biol. 67: 1-10); high activity in egg attributable to maternal expression. Wings of homozygous females and hemizygous males obliquely truncated posteriorly; phenotype varies from wings that are wrinkled and blistered and do not extend beyond the tip of the abdomen to normal, with intermediate phenotypes having wings truncated to various degrees but not wrinkled or blistered, or normal wings with irregularly spaced marginal hairs. Wing cells smaller than normal (Fausto-Sterling and Hsieh, 1975, Dev. Biol. 51: 269-81); oblique truncation attributed to cell death in distal portion of presumptive wing blade (Fausto- Sterling, 1980, J. Exp. Zool. 213: 383-90). Homozygous females usually sterile when crossed to r male; occasionally give a few offspring, virtually all daughters plus a few exceptional males, when outrcossed. r39 females produce many malformed eggs and unfertilized eggs with normal morphology; ovarian development often retarded or fails; yolk deposition affected; lethal effect in progeny results from generalized disturbance in differentiation 13-16 hr after fer- tilization at 25; surviving embryos hatch late and may produce larvae that neither move nor feed (Counce, 1956, Z. Indukt. Abstamm. Vererbungsl. 87: 482-92). Eggs produced by r9 parents fail to hatch, but can be rescued by the injection of preblastoderm eggs with cytoplasm from either fertilized or unfertilized wild-type eggs or with pyrimidine nucleosides (Okada, Kleinman, and Schneiderman, 1974, Dev. Biol. 37: 55- 62). r/0 tissue in gynandromorphs produced by r/r females confined to abdomen; no such constraint when produced by r/+ females (Fausto-Sterling, 1971, Dev. Biol. 26: 452-63). Mosaic studies of Falk (1977, Dev. Biol. 58: 134-47) indicate nonautonomy of r+ within the wing and the ovary, and that nor- mal wing and ovarian development depend on pyrimidine syn- thesis within those organs. Female-sterile but not truncated-wing aspect of phenotype partially alleviated by administration of cytidine during development (Bahn, 1970, DIS 45: 99); administration of 6-azauracil or 6-azauridine, com- petetive inhibitors of pyrimidine synthesis, causes rudimen- tary phenocopies [Rizki and Rizki, 1965, Science 150: 222-23; Str/man, Bahn, N/rby, and Sick, 1973, Hereditas 73: 239-46 (fig.)]. Sex-linked recessive mutants selected on the basis of ina- bility to survive on medium deficient in pyrimidines all map to the rudimentary locus; the majority have the rudimentary phenotype, but some are phenotypcially normal and are desig- nated subliminal alleles in the following table. Subliminal alleles exhibit strongly depressed survival on pyrimidine-free medium; standard alleles do not survive in the absence of pyrimidine; relative survival of r/+ heterozygotes varies from 5 to 70% depending on severity of allele (Falk and Nash, 1974, Mol. Gen. Genet. 131: 339-49). alleles: complementation C????BA A?B??CD allele origin discoverer synonym ( ref | 1234567 / comments ` ___________________________________________________________________________________________________________________________________________ *r1 Morgan, 10f 25, 26, 27 *r2 Bridges, 14g 10 *r2L Lancefield 19 *r3 Sturtevant, 17j30 *r4 spont Wallace *r5 spont Bridges 8 *r6 spont Bridges 8 *r7 spont Bridges 8 *r8 Sturtevant 8 r9 spont Bridges, 20b3 r7, 8 7, 25 +++++-+ *r10 spont Bridges 8 *r11 Sturtevant 8 r12 spont Wallace, 22k8 25 *r13 spont Bridges 8 r13A chemical? Fahmy rMAHI13A, 45 7 ------- *r14 spont Bridges, 24d4 25 *r15 spont Bridges 8 r15 chemical? Fahmy 6 6 ------- *r16 Stern 8 *r17 spont Bridges 8 *r32k Ball 8 *r35 spont Gottschewski, 1935 *r35a X ray Oliver, 35a10 27 *r39 sulfur mustard Auerbach, 1951 11 r39k spont L. V. Morgan, 39k9 rs1, 9 7, 22 +++++-+ r39l spont Buzzati-Traverso, 39l12 6 *r41 sulfur mustard Auerbach, 1951 11 r53l CB. 3025 Fahmy, 53l 29 7, 13 ++-++++ r54c CB. 3026 Fahmy, 54c 36 7, 13 -++++++ r54d CB. 3025 Fahmy, 54d 30 7, 13 --+++++ r54j CB. 3007 Fahmy, 54j 37 7, 13 ------- r55a CB. 3025 Fahmy, 55a 31 7, 13 +--++++ r55k CB. 3034 Fahmy, 55k 19 7, 13 ++++--+ r56d CB. 1528 Fahmy, 56d 25 7, 13 +++---+ r56j CB. 1540 Fahmy, 56j 38 7, 13 ----+++ r56k CB. 1540 Fahmy, 56k 4 7, 13 ++++++- r58a X ray Burdick, 1958 32 7, 5 ----+++ r61 spont Green 18 7 +++++-+ r61i spont Green, 61i8 44 7 ------- r61j spont Green, 61j26 11 7 +++++-+ r62j chemical? Fahmy 22 7 ++----+ r63c spont Clancy, 63c 10 r64k chemical? Fahmy 33 7 ---++++ r68b colchicine Gethmann 32 r68g chemical? Carlson, 68g18 14 7 ++++--- r70b X ray Lefevre, 70b26 29 -?-??-- In(1)7B;15A1-2 r71j spont Mohler, 71j26 29 -?1??+- r75 chemical? Fahmy 7 7 ------- r81k P Osgood, 81k24 28 r1996 chemical? Fahmy 3 7 ++++++- r2291 chemical? Fahmy 23 7 ++++--+ r2381 chemical? Fahmy 10 7 +++++-+ r2622 chemical? Fahmy 16 7 +++++-+ r2696 chemical? Fahmy 35 7 ------- r3433 chemical? Fahmy 34 7 ----+++ r3455 chemical? Fahmy 39 7 --+++++ r3463 chemical? Fahmy 40 7 --+++++ r3484 chemical? Fahmy 5 7 ++++++- r3589 chemical? Fahmy 20 7 ++++--+ r3718 chemical? Fahmy 15 7 +++++-+ r3719 chemical? Fahmy 1 7 ++++++- r3720 chemical? Fahmy 13 7 +++++-+ r3721 chemical? Fahmy 17 7 ++++--+ r3722 chemical? Fahmy 24 7 +------ rbs1 EMS 4 rbs2 EMS 4 rbs3 EMS 4 rC spont N/rby 3, 26 rCS1 EMS Gans 2, 15 cold sensitive lethal rE1 EMS 3 rE2 EMS 3 rE3 EMS 3 rE4 EMS 3 rE5 EMS 3 rE6 EMS 3 rE7 EMS 3 rE8 EMS 3 rE9 EMS 3 rE10 EMS 3 *rG spont Goldschmidt rpx bl 16 *rH H2CO Auerbach 11 rhdt P Tsubota 33 rhd2 P Tsubota 33 rhd3 P Tsubota 33 rhd4 P Tsubota 33 rhd5 P Tsubota 33 *rK Krivshenko 1 In(1)20F rLE1 EMS Rawls 29 +?+??+- rLE2 EMS Rawls 29 +?+??-+ rLE3 EMS Rawls 29 +?+??1+ subliminal allele rLE4 EMS Rawls 29 +?+??-+ rLE5 EMS Rawls 29 +?+??-+ rLE6 EMS Rawls 29 +?+??-+ rLE7 EMS Rawls 29 +?+??-+ rLE8 EMS Rawls 29 -?-??-- rLE9 EMS Rawls 29 +?+??-+ rLE10 EMS Rawls 29 +?+??1+ subliminal allele rLE11 EMS Rawls 29 -?+??++ rLE12 EMS Rawls 29 +?1??-+ rLE13 EMS Rawls 29 +?+??1- rLE14 EMS Rawls 29 -?-??-- rLE15 EMS Rawls 29 +?+??-+ rLE16 EMS Rawls 29 -?-??-- rLE17 EMS Rawls 29 -?-??-- rLE18 EMS Rawls 29 +?+??1+ subliminal allele rLE19 EMS Rawls 29 ++++++- rLE20 EMS Rawls 29 ++++++- rLE21 EMS Rawls 29 +?+??-+ rLE22 EMS Rawls 29 +?+??-+ rLE23 EMS Rawls 29 -?-??-- rLE24 EMS Rawls 29 +?+??-+ rLE25 EMS Rawls 29 -?-??-- rLI1 ICR170 Rawls 29 -?-??-- rLI2 ICR170 Rawls 29 -?-??-- rLI3 ICR170 Rawls 29 -?-??-- rLI4 ICR170 Rawls 29 -?-??-- rLI5 ICR170 Rawls 29 -?-??-- rLI6 ICR170 Rawls 29 -?-??-- rLI7 ICR170 Rawls 29 -?-??-- rLI8 ICR170 Rawls 29 +?+??+- rLI9 ICR170 Rawls 29 -?-??-- rLI10 ICR170 Rawls 29 -?-??-- rLI11 ICR170 Rawls 29 -?-??-- rLI12 ICR170 Rawls 29 -?-??-- rLI13 ICR170 Rawls 29 -?-??-- rLI14 ICR170 Rawls 29 -?-??-- rLI15 ICR170 Rawls 29 -?-??-- rLI16 ICR170 Rawls 29 -?-??-- rLX1 X ray Rawls 29 +?+??-+ rLX2 X ray Rawls 29 +?+??1- rLX3 X ray Rawls 29 -?-??-- rLX4 X ray Rawls 29 +?+??-+ rLX5 X ray Rawls 29 -?-??-- rLX6 X ray Rawls 29 -?-??-- rLX7 X ray Rawls 29 -?-??-- rLX8 X ray Rawls 29 -?-??-- rLX9 X ray Rawls 29 -?-??-- rLX10 X ray Rawls 29 -?-??-- rLX11 X ray Rawls 29 +?-??-+ rLX12 X ray Rawls 29 +?1??-+ rM1 EMS Mohler fs(1)11-722 21 rM2 EMS Mohler fs(1)11-836 21 rM3 EMS Mohler fs(1)11-992 21 rM4 EMS Mohler fs(1)11-007 21 rM5 EMS Mohler fs(1)12-779 21 rM6 EMS Mohler fs(1)12-829 21 rM7 EMS Mohler fs(1)12-1247 21 rM8 EMS Mohler fs(1)12-2088 21 rM9 EMS Mohler fs(1)12-2502 21 rM10 EMS Mohler fs(1)12-3642 21 rM11 EMS Mohler fs(1)12-4331a 21 rM12 EMS Mohler fs(1)13-873 21 rM13 EMS Mohler fs(1)13-1977 21 rM14 EMS Mohler fs(1)14-693 21 +????-+ rM15 EMS Mohler fs(1)14D-73 21 rntg NNG Kaufman 18 rpyr1 EMS 14 -????-- subliminal allele rpyr2 EMS 14 -????-- subliminal allele rpyr3 EMS 14 -????-- temperature-sensitive subliminal allele rpyr4 EMS 14 +????-+ subliminal allele rpyr5 EMS 14 temperature-sensitive subliminal allele rpyr6 EMS 14 +????-+ temperature-sensitive subliminal allele rpyr7 EMS 14 -????-- temperature-sensitive subliminal allele rpyr8 EMS 14 rpyr9 EMS 14 rpyr10 EMS 14 rpyr11 EMS 14 female fertile rpyr12 EMS 14 rpyr13 EMS 14 female fertile rpyr14 EMS 14 +????-+ rpyr15 EMS 14 rpyr16 EMS 14 -????-+ female fertile rpyr17 EMS 14 rpyr18 EMS 14 rpyr19 EMS 14 rpyr20 EMS 14 subliminal allele *rS spont Hadorn, 59d 30 rSch spont Schalet 31 inserted DNA element rs2 spont Green, 59k22 41 7, 17 ----+++ rs3 spont Green, 58b 21 7, 17 ++----+ *rs4 spont Green, 59b 17 *rs5 spont Green, 60i7 17 *rs6 spont Green, 60l12 17 rs7 spont Green, 60l12 17 *rs8 spont Green, 61g2 17 *rSn Silberman 1 rsP1 spont Craymer, 77k 12, 20 rsP2 spont 20 rtsB EMS Baker 12 rX1 X ray Green, 60b13 17 rX2 X ray Green, 60c15 17 rX3 X ray Green, 60c15 28 7, 17 -----++ rX4 X ray Green, 60c15 17 rX5 X ray Green, 60c15 17 rX6 X ray Green, 60dt 42 7, 17 ------- rX7 X ray Green, 60e24 17 rX8 X ray Green, 58a 17 rX9 X ray Gloor, 57a 27 7, 17 +++--++ rX10 X ray Gloor, 57a 26 7, 17 +++--++ rX11 X ray Green, 60k27 43 7, 17 ------- rX12 X ray Green, 60k27 17 rX13 X ray Green, 60k27 12 7, 17 +++++-+ rX14 X ray Green, 62j7 17 rX917 chemical? Fahmy 2 7 ++++++- ( Carlson (1971) ordered 45 alleles by means of genetic recom- bination; the order, from left to right, of alleles thus localized is represented by numbers in bold face. In some publications these numbers are used as allelic designations. | 1 = Agol, 1936, DIS 5: 7; 2 = Azou, Mehl, and Jarry, 1981, Dev. Biol. 84: 157-63; 3 = Bahn, N/rby, and Sick, 1971, Hereditas 69: 187-92; 4 = Bahn and Sondergaard, 1983, Hereditas 99: 309-10; 5 = Burdick, 1961, DIS 35: 45; 6 = Buzzati-Traverso, 1940, DIS 13: 51; 7 = Carlson, 1972, Genet. Res. 19: 129-32; 8 = Carnegie Publication 552; 9 = Carnegie Publication 627; 10 = Clancy, 1964, DIS 39: 65; 11 = Counce, 1956, Z. Indukt. Abstamm. Vererbungsl. 87: 482-92; 12 = Craymer, 1980, DIS 55: 197-200; 13 = Fahmy, 1959, Nature 184: 1927-29; 14 = Falk and Nash, 1974, Mol. Gen. Genet. 131: 339-47; 15 = Gans, Audit, and Masson, 1975, Genetics 81: 683-704; 16 = Goldschmidt, 1945, Univ. Calif. (Berkeley) Publ. Zool. 49: 501-03; 17 = Green, 1963, Genetica 34: 242-53; 18 = Kaufman, 1970, DIS 45: 34; 19 = Lancefield, 1918, Am. Nat. 52: 264-69; 20 = Modolell, Bender, and Meselson, 1983, Proc. Nat. Acad. Sci. USA 80: 1678-82; 21 = Mohler and Carroll, 1984, DIS 60: 236- 41; 22 = Morgan, L.V., 1940, DIS 13: 51; 23 = Morgan, T.H., 1915, Am. Nat. 49: 240-50; 24 = Morgan and Bridges, 1916, Carnegie Inst. Wash. Publ. No. 237: 25 (fig.); 25 = Morgan, Bridges, and Sturtevant, 1925, Bibliog. Genet. 2: 234; 26 = N/rby, 1970, Hereditas 66: 205-14; 27 = Oliver, 1939, DIS 12: 48; 28 = Osgood, 1983, Genetics 104: s55; 29 = Rawls and Porter, 1979, Genetics 93: 143-61; 30 = Rohr, 1962, DIS 36: 39; 31 = Schalet, 1985, Genetics 110: s99; 32 = Thompson, 1969, DIS 44: 44; 33 = Tsubota, Ashburner, and Schedl, 1985, Mol. Cell. Biol. 5: 2567-74. / Complementation results according to Carlson (1972), Rawls and Porter (1979) and Falk (1976, Mol. Gen. Genet. 148: 1- 8; Falk, McCaughin, and Cogley, 1977, Genetics 856: 765- 77); arabic numbers are the equivalent of Roman numerals used by Carlson; upper case letters used by Rawls and Porter, and by Falk, where A of Rawls and Porter corresponds to C of Falk and to 1 of Carlson, B of Rawls and Porter to 3 of Carlson, C of Rawls and Porter to B of Falk to 6 of Carl- son, and D of Rawls and Porter to A of Falk to 7 of Carlson. Lines with seven + or - symbols from Carlson (many confirmed by Rawls and Porter); lines with four + or - symbols from Rawls and Porter only; and lines with but three are from Falk. ` Subliminal alleles exhibit normal-wing phenotype and female fertility, but survive very poorly on pyrimidine deficient medium. Fully mutant derivatives of subliminal alleles induced with EMS by Falk and deBoer (1980, Mol. Gen. Genet. 180: 219-24): six from rpyr1, thirteen from rpyr2, sixteen from rpyr3, 21 from rpyr4, and 29 from rpyr11. - More detailed description below. cytology: Tentatively placed in 15A1 by Lefevre [Genetics and Biology of Drosophila (Ashburner and Novitski, eds.). Academic Press, London, New York, San Francisco, Vol. 1a, pp. 32-66] and in 15A1-2 on the basis of in situ hybridization (Rawls, Freund, Jarry, Louis, Seagraves, and Schedl, 1986, Mol. Gen. Genet. 202: 493-99). molecular biology: r transcription unit selected using a ham- ster pyr1-3 probe; walk in either direction produced 90 kb of cloned sequence from 14F-15A region (Segraves, Louis, Schedl, and Jarry, 1983, Mol. Gen. Genet. 189: 34-40). Genomic probe hybridizes to a 7.3 kb polyadenylated RNA, which is large enough to encode the 220 kd polypeptide. Direction of tran- scription from left to right, beginning with the domain for DHO on the left, continuing through the domain for CPS in the middle and ATC on the right; the region transcribed is approx- imately 12.5 kb in length and contains at least three introns, one of which is over 4 kb in length. other information: Fine structure of the locus examined by both recombination and complementation (Carlson, 1971, Genet. Res. 17: 53-81). Carlson defines seven complementing regions, whereas Fahmy and Fahmy (1959, Nature 184: 1927-29) recog- nized six, Green (1963, Genetica, 34: 242-53) at least three, Falk (1976, Mol. Gen. Genet. 148: 1-8) three, and Rawls and Porter (1979, Genetics 93: 143-61) four. Region D of Rawls and Porter corresponds to regions VII and C of Carlson and Falk respectively, and identifies the DHO domain; region C of Rawls and Porter corresponds to regions VI and B of Carlson and Falk and identifies the CPS domain; and region D of Rawls and Porter corresponds to region I and C of Carlson and Falk and identifies the ACT domain. The genetic map and the com- plementation map are roughly colinear. Noncomplementing alleles map to both the 5' and 3' ends of the gene (Carlson's numbers 5 and 6 at the 5' end and numbers 35, 37, and 42-45 at the 3' end, where numbers refer to approximate linear order of alleles on the genetic map). Complementation maps constructed using wing phenotype and survival on unsupplemented medium not identical (Falk, 1976). # rCS1: rudimentary-cold sensitive phenotype: Homozygotes raised at 16 are lethal, at 20 are rudi- mentary, and at 25 are wild type in phenotype; the same results are observed in rCS1/Df(1)r-D17 and rCS1/r1 heterozy- gotes; r1 is deficient for DHO only. Heterozygotes with r11, which is deficient for CPS, are rudimentary at all tempera- tures, and heterozygotes with r29 and r38, which lack ATC activity, are wild type at all temperatures. Cold-sensitive periods during embryogenesis and during organogenesis in the early pupal stage; larval stages cold stable. Activities of DHO isolated from early embryos cold sensitive; CPS exhibits reduced activity at all temperatures; ATC cold stable (Azou, Mehl, and Jarry, 1981, Dev. Biol. 84: 157-63). # R: Roughened location: 3-1.4. discoverer: E. M. Wallace, 35i. phenotype: Eyes of R/+ rough, have some large dark facets. Pho- toreceptor cell 7 frequently absent (Carthew). Male genitalia frequently rotated and male sometimes sterile; viability about 80% wild type. Homozygote semilethal; wings spread. Thorax short; acrostichal hairs deranged, some missing; eyes small. Homozygous female fertile. RK1. alleles: R2, a hybrid-dysgenesis-induced revertant of R, which is hemizygous lethal (Sliter, Henrich, Tucker and Gilbert, 1989, Genetics 123: 327-36). cytology: Placed in 62B8-12 based on reversion of R by Df(3L)R-G2 = Df(3L)62B2-4;62B11-12 and Df(3L)R-G7 = Df(3L)62B8-9;62F2-5 (Sliter, Henrich, Tucker, and Gilbert, 1989, Genetics 123: 327-36). other information: Recently determined to be an allele of Rap1 (Hariharan). #*R51b origin: Recovered among progeny of female treated as embryo with cold shock. discoverer: Mickey, 51b21. references: 1951, DIS 25: 74. 1951, Genetics 36: 565-66. phenotype: Eyes of heterozygote small, oblong, and rough; facets and eye hairs irregular. Viability good. Homozygote lethal. R51b/R has very small eyes, much fusion of facets, and resembles gl and Gl. RK1. # R3-55.4: see M(GPDH) # R3(+): see T(2;3;4)ci+3 # R26-43: see M(ADH) # r-l: rudimentary-like location: 3-70.4 (based on 240 gl-e recombinants). references: Lastowski and Falk, 1980, Genetics 96: 471-78. Rawls, 1980, Mol. Gen. Genet. 178: 43-49. Rawls, 1981, Mol. Gen. Genet. 184: 174-79. Conner and Rawls, 1982, Biochem Genet. 20: 607-19. phenotype: Encodes a polypeptide with the enzyme activities that catalyze the fifth and sixth steps of de novo pyrimidine synthesis: orotate phosphoribosyl transferase [OPRT (EC. 2.4.2.10)] and orotidylate decarboxylase [ODC (EC. 4.1.1.23)] respectively. Thought to exist as a homodimer (Rawls, 1979, Comp. Biochem. Physiol. 62B: 207-16). The first three enzyme activities encoded by r and the fourth by Dhod. OPRT and ODC activities drastically reduced except that OPRT moderately reduced in r-lK7 and ODC above normal in r-lK6. All mutants accumulate orotic acid, the substrate of ODC. Flies hemizy- gous or homozygous for mutants at this locus resemble r in that they have obliquely truncated wings with deranged margi- nal hairs and virtual female sterility; they differ from r in having faintly mottled eyes and severely impaired viability (viability of hypomorphs moderately reduced). r is epistatic to r-l in that r; r-l flies display the r rather than the r-l phenotype, suggesting that mottled eyes and reduced viability are the consequence of orotic-acid accumulation rather than pyrimidine deficiency. alleles: allele origin discoverer synonym ref ( comments ________________________________________________________________ r-l1 EMS rala1a 2 r-lK1 EMS Rawls 1, 3, 4 noncomplementing r-lK2 EMS Rawls 1, 3, 4 hypomorph; complements r-lK6, r-lK7, and r-lK8 r-lK3 EMS Rawls 1, 4 noncomplementing r-lK4 EMS Rawls 1, 4 noncomplementing r-lK5 EMS Rawls 1, 4 hypomorph; complements r-lK6 and r-lK8 r-lK6 EMS Rawls 1, 4 complements r-lK2, r-lK5, and r-lK7 r-lK7 EMS Rawls 1, 4 complements r-lK2, r-lK6, and r-lK8 r-lK8 EMS Rawls 1, 4 complements r-lK2, r-lK5, and r-lK7 ( 1 = Conner and Rawls, 1982, Biochem. Genet. 20: 607-19; 2 = Lastowski and Falk, 1980, Genetics 96: 471-78; 3 = Rawls, 1980, Mol. Gen. Genet. 178: 43-49; 4 = Rawls, 1981, Mol. Gen. Genet. 184: 174-79. cytology: Tentatively placed in 93B4-13 based on its inclusion in Df(3R)e-R6 = Df(3R)93B4-5;94A5-16 (B.S. Baker) and Df(3R)e-R1 = Df(3R)93B3-5;93D2-4 (B.S. Baker) but not Df(3R)e-F2 = Df(3R)93A6-B1;93D7-10 (Mohler and Pardue) = Df(3R)93B8-13;93F9-10 (Scalenghe and Ritossa). # r((GPDH): regulator of ( glycerol phosphate dehydrogenase location: 3-55.4. origin: Spontaneous. synonym: m(GPDH), r3-55.4. references: King and McDonald, 1982, DIS 58: 91-92. 1983, Genetics 105: 55-69. phenotype: Recessive allele of a locus that affects levels of (glycerol phosphate dehydrogenase activity. In the presence of the dominant allele, which we designate r((GPDH)+, (GPDH levels in the adult are 60% higher than they are in r((GPDH) homozygotes; the increase is observed in the thorax but not the abdomen. Rates of enzyme synthesis the same in the two genotypes; differences in activity attributed to reduced rates of degradation in the presence of r((GPDH)+; no evidence for conformational differences in (GPDH. Not a general regulator of enzyme activity; however one rapidly migrating polypeptide is much more abundant in r((GPDH) homozygotes than in the presence of r((GPDH)+. Larvae carrying r((GPDH)+ have 25% lower levels of (GPDH activity than r((GPDH) homozygotes. # r(ADH) location: 3- (between h and th). origin: Naturally occurring allele. synonym: r26-43. references: King and McDonald, 1987, Genetics 115: 693-99. phenotype: r(ADH) homozygotes exhibit approximately 70% of the level of alcohol dehydrogenase seen in heterozygotes and r(ADH)+ homozygotes; parallel differences in the levels of immunologically cross-reacting material (CRM) also observed. The effect of r(ADH) is constant over development and is not attributable to differences in charge, conformation, or rate of synthesis of ADH. Activities of GPDH and PGI levels insen- sitive to r(ADH) genotype. # ra: rase location: 3-97.3. origin: Spontaneous. discoverer: Beadle, 34d. references: 1935, DIS 4: 10. phenotype: Bristles and hairs small and irregularly absent, especially from head and thorax. Viability good; developmen- tal time normal. RK2. cytology: Associated with In(3R)P (Craymer). #*ra2 origin: Spontaneous in In(3R)P. discoverer: Mossige, 36k21. synonym: bd: bald. references: 1937, DIS 8: 9. phenotype: Homozygote lacks all head bristles and some scutel- lars. Heterozygote has extra anterior scutellars in about 30% of flies. RK2A. cytology: Occurred in and probably inseparable from In(3R)P = In(3R)89C2-4;96A18-19. #*rab: rabbit location: 1-58. origin: Induced by P32. references: Bateman, 1950, DIS 24: 55. phenotype: Hairs on mesonotum near dorsocentral bristles turned inward toward midline. Air bubbles occasionally in thorax, beneath dorsocentrals, and scutellum. Wings rarely held up. Viability and fertility normal. RK2(A). other information: Slight disturbance of crossing over proximally. # rad201: radiation sensitive location: 2-59.9. synonym: rad(2)201G1. references: Levina, Malinovsky, and Zakharov, 1980, Genetika (Moscow) 16: 285-89. Khromykh and Zakharov, 1981, Genetika (Moscow) 17: 658-66. Khromykh, 1981, Genetika (Moscow) 17: 667-76. Tikhomirova and Rasheva, 1983, Genetika (Moscow) 19: 628-34. phenotype: Survival of homozygous larvae more sensitive to exposure to / radiation than controls; chromosome- and chromatid-aberration induction increased over control but ratio of the two types same as control. X-ray-induced X- chromosome loss in immature oocytes greater in rad201 than in control; mature oocytes show no effect of rad201. Post- treatment hyperthermia enhances the effect of irradiation in mature oocytes and control immature oocytes, but not in imma- ture oocytes of rad201. # radioresistant: see rar1 # radius incompletus: see ri # raf: see phl # rag: rags (T. Schupbach) location: 2-32. origin: Induced by ethyl methanesulfonate. references: Schupbach and Wieschaus, 1989, Genetics 121: 101- 17. phenotype: Maternal effect lethal; embryos from homozygous mothers form a fragmented cuticle with large ventral holes and head defects. alleles: One: ragQI = rag1. #*rag: ragged location: 3-37 (Steinberg). discoverer: Charles, 1932. references: Dunn, 1934, DIS 1: 30. phenotype: Hairs missing from sections of wing margin. RK3. # rags: see rag #*rai: raisin location: 3-17 (Stanley). origin: Spontaneous. discoverer: Hersh. references: 1953, DIS 27: 55. phenotype: Eye color deep brown, like se. Eclosion delayed 1 or 2 days. RK2. # raised: see rsd # raised wing: see rw # raisin: see rai # ral: see r-l # rap: retina aberrant in pattern (J.C. Hall) location: 1- {7.5}. references: Karpilov, Kolodkin, Bork, and Venkatesh, 1989, Genes Dev. 3: 1834-44. phenotype: Eyes grossly abnormal: rough surface texture, and deep pseudopupil (DPP) has anomalous appearance; finer level observations show ommatidia to be aberrant in size, shape, and alignment; in sections, numbers of photoreceptor cells per ommatidium vary greatly, and positions of such cells plus rhabdomere arrangements are altered. Developmentally, recruitment of presumptive photoreceptors behind morphogenetic furrow in third larval instar eye disc is abnormal: e.g., arrangement of five-cell preclusters is very disorganized, and the normal increase in numbers of cells within clusters does not occur, so that, in regions where eight-cell clusters are found in wild-type, there are small clusters in rap, intermin- gled with others than can have >8 cells; cell bodies of developing photoreceptor cells do not become apically posi- tioned in the disc, as in wild-type. Temperature-shift experiments on rap4 show a temperature-sensitive period for roughened eye is in third larval instar. Mosaic experiments, involving induced mitotic recombination in rap1/+ larvae, showed that normal adult ommatidia always had a rap+ R8 cell, whereas R1-7 could readily be genotypically rap1 in such facets; also, certain phenotypically mutant ommatidia, near borders separating mutant and wild-type tissue, contained all rap+ cells (in clusters of 3, 4, 6, or 8 photoreceptor cells). alleles: allele origin discoverer synonym comments _______________________________________________________________ rap1 P hypomorph? rap2 P hypomorph? rap3 P amorph rap4 P rapR22ts temperature-sensitive hypomorph ( ( Rearing at 17 leads to normal eye-surface morphology (albeit 36% penetrance of rap1-like DPP phenotype); rearing at 29 causes all flies to have relatively mild eye roughening. cytology: Maps to 4C7-16, based on its inclusion in Df(1)rb13 = Df(1)4C5-6;4D3-E1 but not Df(1)rb1 = Df(1)3F6-4A1;4C7-8 or Df(1)JC70 = Df(1)4C15-16;5A1-2. other information: rap1 was shown to complement the nearby rough-eye mutation, rg. # Rap1: Ras3 location: 3- 1.4 (based on mapping of R). origin: Isolated from genome library using v-Ha-ras probe. synonym: Dras3, Ras3. references: Neuman-Silberberg, Scheifer, Hoffmann, and Shilo, 1984, Cell 37: 1027-33. cytology: Cloned sequence hybridizes to 62B. molecular biology: Transcripts of 1.5, 1.9, and 2.9 kb detected (Lev, Kimchie, Hessel, and Segev, 1985, Mol. Cell. Biol. 5: 1540-1542); larger transcript present throughout develop- ment; shorter ones more abundant during embryogenesis. other information: Isolated as a Ras homologue, but recently found to share high homology with human Rap. # rapid exhaustion: see rex # rar1: radioresistant location: 1- (not mapped). origin: Spontaneous in line selected for radioresistance. references: Nothel, 1980, DIS 55: 208-09. Nothel, 1981, Mutat. Res. 80: 105-20. Nothel, 1982, Mutat. Res. 103: 87-90. Nothel and Abdalla, 1982, Mutat. Res. 92: 123-32. phenotype: Semidominant; reduces sensitivity of prestage-14 vitellarial oocytes to the dominant- and recessive-lethal- inducting effects of X irradiation or exposure to ethyl methanesulfonate. 1.3 X the dose to produce a given effect in controls required to produce the same effect in rad1 (i.e., Dose Reduction Factor = 1.31). Effect of mutation inhibited by caffein. # rar2 location: 2- (near centromere). origin: Spontaneous in line selected for radioresistance. references: Nothel, 1980, DIS 55: 208-09. Nothel, 1981, Mutat. Res. 80: 105-20. Nothel, 1982, Mutat. Res. 103: 87-90. phenotype: Semidominant; produces relative resistance of imma- ture oocytes to X-ray and ethyl-methanesulfonate induction of dominant and sex-linked recessive lethals (Dose Reduction Fac- tor = 1.31) and X-chromosome loss (Dose Reduction Factor = 1.72). # rar3 location: 3-49.8. origin: Spontaneous in line selected for radioresistance. references: Nothel, 1980, DIS 55: 208-09. Nothel, 1981, Mutat. Res. 80: 105-20. Nothel, 1981, Mutat. Res. 84: 291-304, 305-13, 315-19. Nothel, 1982, Mutat. Res. 103: 87-90. phenotype: Recessive mutant conferring resistance to the genetic effects of irradiation on oogonia and immature oocytes. Confers a dose reduction factor of 1.58 on the induction of dominant lethals, nondisjunction of major chromo- somes and homologous exchange; and of 1.87 on the induction of sex-linked recessive lethals. Without effect on male germ line or somatic cells. Resistant to ethyl methanesulfonate as well. # ras: raspberry location: 1-32.35 (Lefevre). phenotype: Defective in pteridine synthesis; eye color dark ruby. Appears to affect the level of guanosine triphosphate cyclohydrolase activity; ras alleles exhibit increased activity compared to wild type at pupariation, but decreased activity at eclosion (Evans and Howell, 1979, Biochem. Genet. 16: 13-26). Not a purine auxotroph. Color autonomous in lar- val optic disks transplanted into wild-type hosts (Beadle and Ephrussi, 1936, Genetics 21: 230). ras/rasl display ras eye color. Larval Malpighian tubes nearly wild type; not useful for classification (Brehme and Demerec, 1942, Growth 6: 351- 56). ras is but one of four types of mutations belonging to an interrelated complex involved some way in purine metabol- ism: ras, a nonauxotrophic eye-color mutant; two purine auxo- trophs, pur1 and gua1, which complement ras and one another; and rasl, lethal alleles that fail to complement completely the other three types. The complex shares genetic features with rudimentary. alleles: allele origin discoverer synonym ref ( comments ____________________________________________________________________________ ras1 heat Muller, 28d17 9 ras2 Grossman, 1932 2 ras3 spont Ives, 37b18 eye color lighter than ras1; darkens with age ras4 spont Ives, 38f female sterile ras5 P Green rasf2 3 ras6 P Green rasf26 3 ras7 P Green rasj26 3 ras8 fast Lewis, 1953 rasv 1 variegated; neutron Tp(1;3)9E;13C;81F rasl1 EMS Grigliatti, 1970 l(1)E6ts 4 temperature- sensitive lethal rasl2 X ray Lefevre l(1)HC199 6,7 lethal rasl3 X ray Lefevre l(1)KC12 7 lethal rasl4 X ray Lefevre l(1)RC1 7 lethal rasl5 X ray Lefevre l(1)S27 7 lethal rasl6 X ray Lefevre l(1)S89 7 lethal rasl7 EMS Lefevre l(1)DA589 6,8 lethal rasl8 EMS Lefevre l(1)DC819 6,8 lethal rasl9 EMS Lefevre l(1)DF955 6,8 lethal rasl10 EMS Lefevre l(1)EA140 6,8 lethal rasl11 EMS Lefevre l(1)VA212 6,8 lethal rasl12 EMS Lefevre l(1)VD224 6,8 lethal rasl13 EMS Lefevre l(1)VE432 6,8 lethal rasl14 EMS l(1)D12 5,10 lethal rasl15 EMS l(1)F1 5,10 lethal| rasl16 EMS l(1)F19 5,10 lethal| rasl17 EMS l(1)H6 5,10 lethal rasl18 EMS l(1)H25 5,10 lethal rasl19 EMS l(1)J5 5,10 haplo-specific lethal/ rasl20 EMS l(1)K27 5,10 lethal rasl21 EMS l(1)N23 5,10 lethal ( 1 = Brokaw, 1954, DIS 28: 73; 2 = Dunn, 1934, DIS 1: 30; 3 = Green, 1977, Proc. Nat. Acad. Sci. USA 74: 3490-93; 4 = Grigliatti and Suzuki, 1970, Proc. Nat. Acad. Sci. USA 67: 1101-08; 5 = Janca, Woloshyn, and Nash, 1986, Genetics 112: 43-64; 6 = Johnson, Woloshyn, and Nash, 1979, Mol. Gen. Genet. 174: 287-92; 7 = Lefevre 1981, Genetics 99: 461-80; 8 = Lefevre and Watkins, 1986, Genetics 113: 869-95; 9 = Muller, 1935, DIS 3: 30; 10 = Nash and Janca, 1983, Genetics 105: 957-86. | Complements ras2 eye color but not lethality of other rasl alleles. / Survives weakly as homozygous female, but completely lethal as male and hemizygous female. cytology: Placed in 9E1-4 on the basis of its inclusion in the region of overlap between Df(1)ras-P14 = Df(1)9E1-2;9F3-4 and Dp(1;2)9E1;10A11;56A (Zhimulev, Pokholkova, Bgatov, Semeshin, and Belyaeva, 1981, Chromosoma 82: 25-40). # Ras1: Ras proto-oncogene sequence location: 3- {49}. origin: Isolated from genomic library using v-Ha-ras probe. synonym: Dras1. references: Neuman-Silberberg, Scheifer, Hoffmann, and Shilo, 1984, Cell 37: 1027-33. phenotype: Codes for a polypeptide of 21.6 kilodaltons; resi- dues 1-121 and 137-64 exhibit 75% homology with vertebrate H- ras protein. alleles: Loss-of-function mutations are homozygous lethal; enhance sev and EgfrE in heterozygotes (M. Simon). cytology: Cloned sequence hybridizes to 85D. molecular biology: Gene cloned and sequenced. Contains four introns. Open reading frame in cDNA codes for 189 amino acids corresponding to a polypeptide of 21 kilodaltons. Transcripts of 1.3 and 2.0 kb detected by Lev, Kimchie, Hessel, and Segev (1985, Mol. Cell. Biol. 5: 1540-42); the larger persists throughout development, the smaller being primarily in unfer- tilized eggs and embryos. # Ras2 location: 3- {15}. origin: Isolated from genome library using v-Ha-ras probe. synonym: Dras2, Dmras64B. references: Neuman-Silberberg, Scheifer, Hoffmann, and Shilo, 1984, Cell 37: 1027-33. Moser, Marlor, Parkhurst, and Corces, 1985, Mol. Cell. Biol. 5: 885-89. phenotype: Encodes a 22.7 kilodalton, 195 amino acid, polypep- tide, p21. cytology: Cloned sequence hybridizes to 64B. molecular biology: Entire gene sequenced. Contains two more amino-terminal codons than the homologous gene from humans and four fewer than the yeast gene. Drosophila, human, and yeast sequences highly homologous up to amino-acid residue 90, less so for the next 80 amino acids, and not at all for residues 170-190. Ras2 contains two introns in positions different from those in the homologous human gene. Appears to produce transcripts of 1.6, 2.1, and 2.6 kb, which are present in the same proportions throughout development (Wadsworth, Madhaven, and Bilodeau-Wentworth, 1985, Nucl. Acids Res. 13: 2153-70; Mozer, Marlo, Parkhurst, and Corces, 1985, Cell. Biol. 5: 885-89). Claimed by Lev, Kimchie, Hessel, and Segev (1985, Mol. Cell. Biol. 5: 1540-42) to produce transcripts of 1.4 and 1.8 kb, the larger of which is expressed at all stages and the smaller in unfertilized eggs and early embryos. # Ras3: see Rap1 # Ras proto-oncogene sequence: see Ras1 # rase: see ra # raspberry: see ras # raspberry-lethal: see ras-l # rauhig: see gl3 # raven: see rv # raw: raw location: 2-19. origin: Induced by ethyl methanesulfonate. references: Nusslein-Volhard, Wieschaus, and Kluding, 1984, Wilhelm Roux's Arch. Dev. Biol. 193: 267-82. Tearle and Nusslein-Volhard, 1987, DIS 66: 209-26. phenotype: Homozygous lethal; dorsal closure and cuticle dif- ferentiation defective. alleles: raw1 and raw2 recovered as IG and IIF. # rb: ruby location: 1-7.5. discoverer: Bridges, 14j18. phenotype: Eye color clear ruby, white in combination with wa, orange with st, and brownish red with bw (Mainx, 1938, Z. Indukt. Abstamm. Vererbungsl. 75: 256-76). Development of pigment autonomous in rb eye disks transplanted into wild-type hosts (Beadle and Ephrussi, 1936, Genetics 21: 230). Larval Malpighian tubes pale yellow (Beadle, 1937, Genetics 22: 587-611). RK1. alleles: allele origin discoverer ref ( ________________________________________ rb1 Bridges, 14j18 rb48a X ray Fox, 48a7 2, 3 rb64f X ray Ives, 64f14 rb66a X ray Becker 1 rbm48 X ray 5 rbntg NNG Kaufman 4 ( 1 = Becker, 1968, DIS 43: 59; 2 = Fox, 1948, DIS 22: 53; 3 = Fox, 1949, Genetics 34: 647-64; 4 = Kaufman, 1970, DIS 45: 34; 5 = Valencia, 1966, DIS 41: 58. cytology: Located in 4C6 based on the overlap of Df(1)rb13 = Df(1)4C5-6;4D3-E1 and Df(1)rb46 = Df(1)4A3-6;4C6-7 (Banga, Bloomquist, Brodberg, Pye, Larrivee, Mason, Boyd, and Pak, 1983, Chromosoma 93: 341-46). # rbc: see rc # rbl: reduced bristle location: 2-82.3. origin: Spontaneous; natural population. references: Watanabe, 1969, Jpn. J. Genet. 44: 15-22. Watanabe and Oshima, 1970, Genetics 64: 93-106. phenotype: Homozygotes show reduced dorsal bristles; both sexes fertile, but development delayed by 1.5 days and viability reduced. # rbp: see BRC # rc: red cells location: 2-36.8 (between d and J). origin: Spontaneous. discoverer: E. B. Lewis, 1946. synonym: rbc: red blood cells. references: 1950, DIS 24: 59. Jones and Lewis, 1957, Biol. Bull. 112: 220-24 (fig.). Grell, 1961, Genetics 46: 925-33. phenotype: rc/rc normal; in lys rc/lys rc, fat cells of head and thorax acquire brownish red pigment. Effect most prom- inant in one or more rows of pigmented cells along mid-dorsal line of thorax just beneath chitin. Pigment is ommochrome since lys rc bw cells are pigmented, whereas v; lys rc cells are colorless except in kynurenine-fed flies. RK3. # rc2 origin: Spontaneous. discoverer: R. F. Grell, 1957. references: Grell, 1961, Genetics 46: 925-33. phenotype: Wild type at 25 on standard medium; at 17, a few red fat cells are visible. Early third instar larvae placed on glucose-agar medium produce flies with numerous red cells. lys rc2 has red cells under any conditions. RK3. rd: reduced From Bridges and Brehme, 1944, Carnegie Inst. Washington Publ. No. 552: 157.