# w: white location: 1-1.5. references: Morgan, 1910, Science 32: 120-22. Morgan and Bridges, 1916, Carnegie Inst. Washington Publ. No. 237: 25, 28, 51. Muller, 1932, Proc. Intern. Congr. Genet. 6th, Vol. 1: 234. Muller, 1935, J. Genet. 30: 407-14. Beadle and Ephrussi, 1936, Genetics 21: 230. Sturtevant and Beadle, 1939, An Introduction to Genetics, W.B. Saunders Co., Philadelphia, p.64 (fig). Brehme and Demerec, 1942, Growth 6: 351-56. Hadorn and Mitchell, 1951, Proc. Nat. Acad. Sci. USA 37: 650-65. Lewis, 1952, Proc. Nat. Acad. Sci. USA 38: 953-61. MacKendrick and Pontecorvo, 1952, Experientia 8: 390-91. Lewis, 1956, Genetics 41: 651. Judd, 1957, Genetics 42: 379-80. 1958, Proc. Intern. Congr. Genet., 10th, Vol. 2: 137. Ziegler-Gunder and Hadorn, 1958, Z. Indukt. Abstamm. Vere- bungsl. 89: 235-45. Green, 1959a, Heredity 113: 303-15. 1959b, Proc. Nat. Acad. Sci. USA 45: 549-53. 1959c, Z. Indukt. Abstamm. Vererbungsl. 90: 375-84. Judd, 1959, Genetics 44: 34-42. Sullivan and Sullivan, 1975, Biochem. Genet. 13: 603-13. Green, 1976, The Genetics and Biology of Drosophila (Ashburner and Novitski, eds.). Academic Press, London, New York, San Francisco, Vol. 1b, pp. 929-46. Judd, 1976, The Genetics and Biology of Drosophila (Ashburner and Novitski, eds.). Academic Press, London, New York, San Francisco, Vol. 1b, pp. 767-99. Gehring and Paro, 1980, Cell 19: 897-904. Bingham, 1981, Cold Spring Harbor Symp. Quant. Biol. 45: 519-25. Bingham and Judd, 1981, Cell 25: 705-11. Bingham, Levis, and Rubin, 1981, Cell 25: 693-704. Collins and Rubin, 1982, Cell 30: 71-79. Goldberg, Paro, and Gehring, , 1982, EMBO J. 1: 93-98. Levis, Bingham, and Rubin, 1982, Proc. Nat. Acad. Sci. USA 79: 564-68. Levis and Rubin, 1982, Cell 30: 543-50. Zachar and Bingham, 1982, Cell 30: 529-41. O'Hare, Levis, and Rubin, 1983, Proc. Nat. Acad. Sci. USA 80: 6917-21. Pirrotta, Hadfield, and Pretorius, 1983, EMBO J. 2: 927-34. Fjose, Polito, Weber, and Gehring, 1984, EMBO J. 3: 2087-94. Gehring, Klemenz, Weber, and Kloter, 1984, EMBO J. 3: 2077- 85. Hazelrigg, Levis, and Rubin, 1984, Cell 36: 469-81. Levis, O'Hare, and Rubin, 1984, Cell 38: 471-81. O'Hare, Murphy, Levis, and Rubin, 1984, J. Mol. Biol. 180: 437-55. Pirrotta and Brockl, 1984, EMBO J. 3: 563-68. Carbonara and Gehring, 1985, Mol. Gen. Genet. 199: 1-6. Chapman and Bingham, 1985, DIS 61: 48-50. Davison, Chapman, Wedeen, and Bingham, 1985, Genetics 110: 479-94. Levis, Hazelrigg, and Rubin, 1985, EMBO J. 4: 3487-99. Pirrotta, Steller, and Bozzetti, 1985, EMBO J. 4: 3501-08. Rubin, Hazelrigg, Karess, Laski, Laverty, Levis, Rio, Spencer, and Zuker, 1985, Cold Spring Harbor Symp. Quant. Biol. 50: 529-33. Steller and Pirrotta, 1985, EMBO J. 4: 3765-72. Zachar, Chapman, and Bingham, 1985, Cold Spring Harbor Symp. Quant. Biol. 50: 337-46. Bingham and Chapman, 1986, EMBO J. 5: 3343-51. Davis, Shen, and Judd, 1987, Proc. Nat. Acad. Sci. USA 84: 174-78. Hazelrigg, 1987, TIG 3: 43-47. Judd, 1987, Structure and Function of Eukaryotic Chromosomes (Hennig, ed.). Springer-Verlag, Berlin, Heidelberg, pp. 81- 94. Mount, 1987, Nature (London) 325: 487. Pastink, Schalet, Vreeken, Paradi, and Eeken, 1987, Mutat. Res. 177: 101-15. Dreesen, Johnson, and Henikoff, 1988, Mol. Cell Biol. 8: 5206-15. Miyashita and Langley, 1988, Genetics 120: 119-212. Mount, Green, and Rubin, 1988, Genetics 118: 221-34. Pastink, Vreeken, and Vogel, 1988, Mutat. Res. 199: 47-53. Birchler and Hiebert, 1989, Genetics 122: 129-38. Birchler, Hiebert, and Rabinow, 1989, Genes Dev. 3: 73-84. Rabinow and Birchler, 1989, EMBO J. 8: 879-89. Tearle, Belote, McKeown, Baker, and Howells, 1989, Genetics 122: 595-606. Gubb, Ashburner, Roote, and Davis, 1990, Genetics 126: 167- 76. phenotype: The white locus is involved in the production and distribution of ommochrome (brown) and pteridine (red) pig- ments found in the compound eyes and ocelli of adult flies as well as the pigments in adult testis sheaths and larval Mal- pighian tubules; the specific function of the protein it encodes is still unknown, but it is believed to be a membrane-associated ATP-binding transport protein for pigment precursors in both the ommochrome and pteridine pathways (Sul- livan and Sullivan, 1975; Mount, 1987; Dreesen et al., 1988; Tearle et al., 1989). w1 was the first mutant found in Droso- phila melanogaster (Morgan, 1910; Morgan and Bridges, 1916). Mutant alleles do not appreciably affect the viability and fertility of the flies. Extreme white alleles as well as white deficiencies remove both brown and red pigments, the w1 allele having very little, if any, pteridine (Hadorn and Mitchell, 1951); isoxanthopterin is present in considerable quantity during pupation but is eliminated during the first three days of adult life (Hadorn, 1954, Experientia 10: 483- 84). Hypomorphic alleles are visibly lighter in combination with w1 than when present as homozygotes. Intermediate white alleles result in partial loss of ommochromes and pteridines; some alleles also affect the distribution of these pigments in the compound eyes (Lewis, 1956; Green, 1959a, 1959c). Although the mutants are positively phototactic, they show no optomotor responses (Kalmus, 1943, J. Genet. 45: 206-13). Wild-type alleles are incompletely dominant over mutant alleles, w/w+ heterozygotes, though visibly indistinguishable from w+/w+, have less red pigment (Muller, 1935; Ziegler- Gunder and Hadorn, 1958; Green, 1959b). Mutant larval disks transplanted into wild-type host develop autonomously (Beadle and Ephrussi, 1936). Early genetic studies identified mutations separable by intralocus recombination into at least seven groups spanning 0.03 cm (Lewis, 1952; MacKendrick and Pontecorvo, 1952; Green, 1959a; Judd, 1959). Mutants occupying the centromere-proximal sites apparently play a regulatory role (Judd, 1976). Subse- quent molecular analysis has localized the proximal mutations to the 5' end of the transcription unit (we) and the upstream flanking sequences (wsp) (Judd, 1987). Mutations at the distal sites have been mapped to the protein coding exons and the introns between them. The proximally-located regulatory mutants (we, for example) do not show dosage compensation; they suppress the zeste gene, and some of them (the wsp alleles) affect the distribution of the red and brown screen- ing pigments of the eyes. Most of the distally-located structural mutants show dosage compensation, wa/Y males having the same eye color as wa/wa females, and do not suppress (but may interact with) zeste. Green (1959a) found that wi fails to show dosage compensation and does not suppress zeste; but wh exhi- bits both zeste suppression and dosage compensation. In spite of their heterogeneity, the alleles at the white locus fail to complement each other except for wsp which partially comple- ments all other w alleles except in the presence of za [Babu and Bhat, 1980, Development and Neurobiology of Drosophila, (Siddiqi, Babu, Hall, and Hall, eds.). Plenum Press, New York and London, pp. 35-40)]. Some white alleles (wc for example) are extremely unstable (Green, 1976); w1 is slightly unstable, giving rise to we and wh, mutants with darker eyes than w1. The locus is characterized by asymmetrical recombination involving transposons; the mutants wr,def and wr,dup are the result of such exchange (Davis et al., 1987). Some P-element white transformations show reproducible patterns of pigmenta- tion which can be altered by the trans-acting gene zeste (Rubin et al., 1985). alleles: Mutant and wild-type w alleles are tabulated below. Deficiencies are described in the rearrangement section. The first table includes alleles believed to be extant, the second alleles known to be lost. See end of text for more detailed description of certain alleles. Table I synonym/ superscript allele origin discov. name ref ( comments cytology _________________________________________________________________________________________________________________________________________________ w1 spont Morgan, 10c 10, 46, 54, 56, eyes white; ocelli, adult 67, 83, 96, testis sheath, larval 112, 116, 119, Malpighian tubes colorless; 125, 130-31, suppresses z; com- 140 plements wsp | w3A3 X ray Paradi 153a partially complements wsp / w5 X ray Green 59 like w1; mutates to (from wc) w+, wdc, wi w5a neutrons Schalet 153a eyes pigmented; partially T(1;3)3C;77 + complements wsp / T(1;3)1A;80 + In(3L)67;77 w8 X ray Green 59, 66a, 162 like w1; mutates to (from wc) w+, wc, wdc, wdi w8x1 spont Welshons 66a, 183 reinversion of In(1)N66h26 abnormality in 3C1 w8x2 spont Welshons 66a, 183 reinversion of In(1)N66h26 appears normal w9 X ray Green 59 like w1; mutates to (from wc) w+, we, wi w9A1 X ray Paradi 153a semilethal; partially coding region complements wsp / deletion w10gA / ray Alexandrov 1-2, 4 like w1; suppresses z w11E4 X ray Gans 46 eyes white; suppresses z w11G3 see z11G3 w13D1 X ray Paradi 153a, 153b slowly complements wsp ` coding region deletion w13gA / ray Alexandrov 1, 4 does not suppress z w15gA / ray Alexandrov 1, 4 does not suppress z w17D2 X ray Paradi 153a, 153b eyes pigmented; coding region complements wsp | deletion w17g Kalisch 93 like w1; does not suppress z w20C2 X ray Pastink 153a slowly complements wsp ` w21D2 X ray Pastink 153a partially complements wsp / w25B2 X ray Pastink 153a slowly complements wsp ` w30 see we2 w30C neutrons Schalet 153a does not complement wsp - T(1;3) w31D1 X ray Paradi 153a slowly complements wsp ` T(1;3)3C;64B-C w32k see wbf2 w33e31 see wdil w33l see wsat w35A2 X ray Paradi 153a, 153b slowly complements wsp ` coding region deletion w40aH1 X ray Valencia 182 eyes white; male lethal In(1)1A1-C3;4C4-7; 17B7-8;18E2-3 w41C1 X ray Paradi 153a, 153b slowly complements wsp ` coding region deletion w49 neutrons Schalet 153a does not complement wsp - T(1;2) w51A neutrons Schalet 153a slowly complements wsp ` w53A neutrons Schalet 153a does not complement wsp - Dp(1;1)3A3;3C1 w54 neutrons Schalet 153a partially complements wsp / w57gA / ray Alexandrov 1, 4 does not suppress z w59gA / ray Alexandrov 1, 4 does not suppress z w60 spont Hollander, 75 like w1 (from wa) 1960 w60gA / ray Alexandrov 1 does not suppress z w61a spont Hess 72 eyes white at 24; yellowish brown at 28 w62k spont 19 w+ revertant (from wi) stable (unlike wi) w64g3 spont Kidd, 1964 102 eyes dark carnation w65a25 X ray Lefevre 15, 100, w65a25/Y eyes white; 113, 176 w65a25/wsp eyes brown; does not suppress z w66A neutrons Schalet 153a does not complement wsp - w66g see w10gA w67a X ray Lefevre 3 eyes white; does not suppress z w67g X ray Lefevre 3 eyes white; does not suppress z w68e EMS Maddern, 68e4 70 eyes white; ocelli white w68g EMS Maddern, 68g13 70 eyes white; ocelli white w68h EMS Maddern, 68h7 70 eyes white; ocelli white w69 NNG Kaufman 99 like w1 w69e spont Neeley, 69c 144 eyes white; ocelli colorless w69gA / ray Alexandrov 1-2, 4 does not suppress z w70C2 X ray Paradi 153a eyes pigmented; complements wsp | w71e spont Whitney, 71e 185 like w1 w73A1 X ray Paradi 153a, 153b slowly complements wsp ` coding region deletion w73d spont Periquet, 73d 5 like w1 w74b / ray Alexandrov 3 eyes ecru; does not suppress z w74d50 / ray Alexandrov 3 eyes white; does not suppress z w74d145 / ray Alexandrov 3 eyes white; does not suppress z w74g EMS Craymer, 74g7 32 like w1 w74k / ray Alexandrov 3 does not suppress z w79b6 / ray Alexandrov 3 eyes white; does not suppress z w80B2 X ray Paradi 153a slowly complements wsp ` w80j1 wc FB Collins, Rubin 30 eyes white elements w80k1 wc FB Collins, Rubin 30 eyes white elements w81d spont Najera 143 eyes white w81e1 wc FB Collins, Rubin 30 eyes white; mutable elements w82a3 wc FB Collins, Rubin 30 eyes white; stable elements w83B X ray Paradi 153a partially complements wsp / T(1;3)3C;64B-C w86.1 transposon- Judd 34, 100 eyes white; mediated does not suppress z deficiency w86.3 transposon- Judd 34, 100 eyes white; mediated does not suppress z duplication w86.5 transposon- Judd 34, 100 eyes white; mediated does not suppress z deficiency w101 ENU Pastink 153c eyes pigmented w102 ENU Pastink 153c eyes pigmented w103 ENU Pastink 153c eyes pigmented w106 ENU Pastink 153c eyes white w107 ENU Pastink 153c eyes pigmented w108 ENU Pastink 153c eyes white w109 ENU Pastink 153c eyes white w113 ENU Pastink 153c eyes white w114 ENU Pastink 153c eyes white w115 ENU Pastink 153c eyes white w116 ENU Pastink 153c eyes pigmented w117 ENU Pastink 153c eyes white w118 ENU Pastink 153c eyes white w119 ENU Pastink 153c eyes white w120 ENU Pastink 153c eyes white w121 ENU Pastink 153c eyes pigmented w123 ENU Pastink 153c eyes pigmented w123D X ray Paradi 153c semi-dominant; In(1)1A;3C; wing deltas; partially Tp(3;1)12F;64B-C;65F complements wsp / w124 ENU Pastink 153c eyes white w126 ENU Pastink 153c eyes pigmented w127 ENU Pastink 153c eyes pigmented w128 ENU Pastink 153c eyes white w130 ENU Pastink 153c eyes pigmented w132 ENU Pastink 153c eyes pigmented w133 ENU Pastink 153c eyes pigmented w133C2 X ray Paradi 153a eyes pigmented; complements wsp w134 ENU Pastink 153c eyes pigmented w136 ENU Pastink 153c eyes white w137 ENU Pastink 153c eyes pigmented w138 ENU Pastink 153c eyes white w141 ENU Pastink 153c eyes pigmented w143 ENU Pastink 153c eyes white w144 ENU Pastink 153c eyes pigmented w147 ENU Pastink 153c eyes white w148 ENU Pastink 153c eyes pigmented w159 ENU Pastink 153c eyes white w163 ENU Pastink 153c eyes white w168 ENU Pastink 153c eyes white w191E neutrons Schalet 153a slowly complements wsp ` w258-12 X ray Demerec, 33j like w1 w781 TE? Valade 181 eyes white; stable del Rio w1118 spont R. Levis 71, 116 eyes white; deletion of (from part of gene wDZL) w#6 HD Simmons, 147, 165, 166, eyes white; Lim 171 mutates to w+ w#12 HD Simmons, 147, 165, 171 eyes white; Lim mutates to w+ w+A Timofeeff- 140, 180 eyes of w/w/w+A Ressovsky triploids pinkish, later maroon w+C Green 55, 56, 58 eyes of w/w/w+C triploids reddish w+O Green 55, 56, 58 eyes of w/w/w+O triploids maroon w+R Timofeeff- 140, 180 eyes of w/w/w+R Ressovsky pinkish, later normal red w+TE Rasmuson eyes w+; unstable w+u spont Gethmann 48, 62, eyes w+; 63 mutates to w- w-r see Df(1)wrJ1 w-r see Df(1)wrJ2 w-rN see Df(1)wrJ3 wa spont Huestis, 1923 apr 10, 17, 17a, 17b, eyes of male yellow-orange, 23, 53-54, 58, of female lighter, yellower; 63, 91, 93, 113, variegated with mw; 126, 132, 135 larval Malpighian tubes 136, 139, 153a, colorless; suppressed by 187, 189 su(wa), lightened by E(wa); does not suppress z; complements wsp |; carries copia wa2 spont Bridges, 1929 24, 25a, 54 eyes orange, darker in male; larval Malpighian tubes colorless; does not suppress z; not affected by su(wa) wa3 spont Curry, 34g2 24, 33, 54 eyes brownish orange; larval Malpighian tubes colorless; does not suppress z; not affected by su(wa) wa3 see wh wa4 spont Nichols-Skoog, 17, 17a, 17b, 24, eyes of male yellow- 35c12 54, 117 orange; of female lighter, yellower (both paler than wa); carries copia; variegated with mw; larval Malpighian tubes colorless; does not suppress z; not affected by su(wa), but affected by E(wa) wa59k13 spont Green, 59k13 158-59 eye color between wa (from wa) and w+; more brown pigment; enhanced by su(f); does not suppress z wa79i spont Najera 143 like wa wa+1 EMS Banerjee 9 revertant of wa wa+2 EMS Banerjee 9 revertant of wa waE see we waLTR1 HD Zachar 189 revertant of wa waM spont Mossige waRM 17a, 17b, 133 eye color between wa and w+; more brown pigment; variegated with mw; suppressed by su(wa); does not suppress z; lightened by E(wa); insertion into copia 5' LTR wapl HD Birchler 17, 17a, 17b insertion mutant; revertant of w1 waR57j see *wa57i waR59k1 X ray Green 27, 135-36 partial revertant of wa; waRsLTR derivative waR79l27 spont 135-36 partial revertant of wa; insertion into copia waR84e7 spont Mossige 135-36 partial revertant of wa waR84e19 spont 135-36 partial revertant of wa waR84h spont Mossige 17a, 17b, 135-36 strong revertant of wa; variegated with mw; affected by su(f) and E(wa) but not su(wa) waRM see waM waS spont Neeley, 68g 144 eyes light orange; ocelli colorless wbf spont Safir, 15g28 buff 17, 24, 34, eyes light buff; somewhat 54, 83, lighter in males; lighter 113, 117, 147, at 19 than at 25; larval 163, 169, 187 Malpighian tubes colorless wbf2 X ray Oliver, 32k16 w32k 24, 54, 148 eyes light buff but darker than wbf or wbf3; larval Malpighian tubes yellow; does not suppress z; carries roo wbf3 spont Curry, 36k9 24 eyes buff; larval Malpighian tubes pale yellow wbl spont Hyde, 14g10 blood 15a, 24, 39, eyes yellowish-ruby, 54, 58, later sepialike; female 77, 82, lighter than male; testis 125, 155 sheaths colorless; larval Malpighian tubes pale yellow at 25; temp. sens.; does not suppress z wbw71g spont Whitney, 71g brownish 185 eyes dark orange over w71e; variegated over T(1;3)wvco; male lethal wBwx spont Mossige, 52a Brownex 81, 83, like bw; wBwx/+ duller, 113, 133, darker than w+; testis 155, 187 sheaths colorless; larval Malpighian tubes pale yellow; does not suppress z wc X ray Green crimson 29-30, eyes crimson; wc mutates Tp(1;3)wc (from wi) 48, 59, to w1, w5, 60-63, w8, w9, w-, wdc, 69, 115, 149, wdi, wi, w+; 162 complements wsp wcf X ray Nicoletti, coffee 145, 184 eyes deep ruby, later 1960 sepialike; larval Malpighian tubes bright yellow wcf2 NNG Kaufman 98 eyes ruby red, darkening to sepia in both males and females; with v, eyes orange; ocelli purplish wch spont Safir, 12j cherry 48, 54, eyes translucent pink; 131, 155, male lighter than female; 168, 187 ocelli pale; adult testis sheaths, larval Malpighian tubes colorless; suppresses z wco spont Lancefield, coral 24, 54, eyes of male deep ruby, 1917 104, 125, later garnetlike; female 126, 155 eyes lighter; larval Malpighian tubes pale yellow; does not suppress z wco2 EMS Farmer 44-45 like wco; suppressed by bw+ wco61 spont Hollander, 76 like wco (from w1) 1961 wcol X ray Demerec, w258-8 54, 105, eyes brick-red to dull, 33j6 colored 155 brownish in young males, lighter in females; does not suppress z wcol+1 EMS Banerjee 9 revertant of wcol wcrr spont Judd, 1962 carrot 86, 113 eyes reddish brown; 155, 187 no sex difference in color; does not suppress z wcrr2 NNG Kaufman 99 eyes orange, darkening to reddish brown in both males and females; ocelli colorless wdc spont Green dark crimson 59, 63, eyes dark crimson; (from wc) 162 mutates to w+, w1, wc, wdc, wi wdil spont Green dilute 59, 63 eye color between (from wc) wc and wi; mutates to w+, w1, wc, wdc, wi wdp chloroethyl Auerbach, deep purple 6, 51, eyes deep purple; methane- 1957 54, 155 no sex difference in color; sulfonate does not suppress z wdt spont Neeley, 68k dirty 144 eyes sooty white; wdt/w grayer than w/w; ocelli colorless wDZL spont Bingham Dominant- 11, 15, dominant over w+; zeste-like 16, 30, eyes dull red brown 114-15, in z+ males, 118, 147, 166, yellow in z males 187, 188 and in homo- and hetero- zygous females (z+ or z) when X's synapsed we spont Morgan, 11h waE; 24, 54, 58 eyes of female yellowish (from w1) eosin 117, 123, pink; eyes of male and we/w 131, 139, female lighter; adult testis 155, 177, 187 sheaths and larval Malpighian tubes colorless; enhanced by e(we); suppresses z we2 spont Hefner, 1925 w30 24, 54, 187 similar to we but darker (from w1) and with less sex difference; larval Malpighian tubes colorless; enhanced by e(we); suppresses z we3 spont Nolte, 1953 146, 187 eyes of female darker (extracted than we female, of from male pinker than we male werb) we59 153a slowly complements wsp ` we+1 EMS Banerjee 9 revertant of we we+2 EMS Banerjee 9 revertant of we wec3 X ray Muller ecru 54 eyes almost white wFM6 spont Kidd, 1964 First 102 eyes white (in FM6) multiple 6 wgd spont Rayle, 1964 garnetoid 161 eyes yellowish ruby, eyes brown in wgd su(f) males at 25; suppresses z wh spont Dunn, 34j27 wa3; 17, 17a, 38, eye color between wbf and (from w1) honey 54, 91, 147, wa4; variegated with mw; 155 larval Malpighian tubes colorless; enhanced by e(we); suppresses z wh82 spont Kennison, 1982 like wh in phenotype (from w1) whd80k17 HD Kidwell hybrid 117, 147, 165 eyes white; reverts dysgenesis to w+ on excision of P-element whd81b6 HD Kidwell 165 eyes white; stable in P and M cytotypes; copia insertion whd81b9 HD Kidwell 147, 165 eyes white; reverts to w+ on excision of P-element whd81b11 HD Kidwell 147, 165, 189 eyes white; stable in P and M cytotypes; copia insertion whd81b25 HD Kidwell 165 eyes white; stable in P and M cytotypes; copia insertion whd81c2 HD Kidwell 165 eyes white; reverts to w+ on excision of P-element wi spont Sturtevant, ivory 20-22, 24, 29, eyes light buff or yellowish, (from w1) 1918 54, 59, 91, 97, lighter in male; larval 121, 123, 137, Malpighian tubes colorless; 147, 149, 153a, does not suppress z; 155, 166 slowly complements wsp ` wi16 Fahmy, 42, 43 like wi; unstable Fahmy wi+A X ray Karess, 97, 147 red eyes; carries 3 kb (from wi) Rubin transposable element F wip spont Bowman, 22, 97 eyes light orange (from wi) Green (partial revertant of w1) wip2 Bowman, 22 partial reversion Green of wi wIR1 HD Pelisson Inducer Reactive 25b, 154, 170 brownish eyes at 20, lighter at 25; enhances z; z1 wIR1 males have mottled orange eyes; carries I factor wIR2 HD Pelisson 154, 170 brown eyes at all temp; carries I factor wIR3 HD Pelisson 25b, 154, 170 brown eyes; carries I factor wIR4 HD Pelisson 154, 170 red-brown eyes at 20; lighter at 25; carries I factor wIR5 HD Pelisson 154, 170 brown eyes at all temp; carries I factor wIR6 HD Pelisson 154, 170 red-brown eyes at all temp; carries I factor wIR7 HD Pelisson 154, 170 eyes white molecular deletion wIR8 HD Pelisson 154, 170 eyes white molecular deletion wis spont Rasmuson isoxanthopterinless 157 eye color normal; male has may be intralocus greatly reduced amount of duplication isoxanthopterin in abdomen; z wis male has variegated eye, between z and w+ in color; probably insert of foreign DNA (R. Jones) wm4 X ray Muller, 1929 mottled 138, 164, 179 eyes mottled; viable and In(1)3C1-2;20F fertile in both sexes (Lefevre) wm4h Reuter, Wolff 164 eyes strongly variegated; In(1)3C1-2;20E-F homozygous lethal wm4rv5 X ray Tartof 179 revertant of wm4 In(1)3C;4A;20 wm4rv6 X ray Tartof 179 revertant of wm4 In(1)1F;3C1-2;20 wm4rv26 X ray Tartof 179 revertant of wm4 In(1)3C;6B;20 wm5 X ray Muller, 1929 18, 138 eyes mottled; T(1;4)3C3-4; viable and 101F1-2 fertile in both sexes wm49a X ray Lefevre, 49a7 wmSp 106-07, 160 eyes have red sectors Tp(1;3)3A10-B1; on white backgound; 3E2-3;80 extra Y -> w+ eye wm51b X ray W.K. Baker, 47 eyes mottled for In(1)3C1-2;20 51b19 w and rst wm51c X ray Lefevre, 107-08 eyes of wm51c/w Tp(1;4)3C1-2; (in wm4) 51c20 variegated, male lethal 3C4-7;20A;101 wm54l neutrons Mickey, 54l3 127 like wm4 In(1)3C3-5;20F wm258-18 X ray Demerec, 33k 37 eyes cream colored, mottled; T(1;4)3C4-5;101 viable, fertile in both sexes wm258-21 X ray Demerec, wVD3 80 eyes, larval Malpighian tubes T(1;4)3E5-6;101F 1934 mottled; mottling more extreme at low temperature; heterozygous female Notch; nearly lethal wm264-58 X ray Demerec, 38d 7-8, 36, eyes variegated; 3 lines associated with induced 73, 172-74 differing in degree of Tp(1;3)N264-58 = (with variegation and parental Tp(1;3)3B2-3; N264-58) effect; homozygous viable in 3D6-7;80D-F female only; maternal effect on variegation wm609e X ray Patterson 64-66 eyes variegated T(1;3)3C2-3; 100C3-4 wm4000 X ray Buzzati- 26 eyes cream colored, darker rearrangement with Traverso, in males; variegated for w, break in w 41l7 rst, fa, co; viable and fertile in both sexes wma see Df(1)N-63b wmD3 Dubinin 167 eyes variegated T(1;4)3C;101 wmJ X ray Jonsson, 110, 112, 179 eyes mottled red and white T(1;4)3C2-3; 61i28 20;102C wmMc X ray McLean 142, 179 eye color light mottled; In(1)3C1-2;20A-C variegated for rst wmo spont Hanly 68, 186 eyes light mottled orange, later darkening; drosopterins reduced to 10% normal; wmo/w -> dark brown eyes; viability good wmSp see wm49a wMt spont Morata wM; Morata 101 eyes white wntg NNG Kaufman nitrosoguanidine 99 like wcrr2 wr spont Muller, 1944 reddish 135-136, wr/wr nearly w+; (wa 141 wr/wa dark maroon; partly believed to be reverted) waLTR derivative wr,def intralocus Judd, 1961 wrdf; recombinant- 34, 84, 86-87, eyes white; wrdf/wsp intralocus exchange deficiency 91 like wsp/wsp; reduces deficiency recombination in w region; for wch site suppresses z wr,dup intralocus Judd, 1961 wrdp; recombinant- 34, 79, 84, eye color lighter than wbf; intralocus exchange duplication 86-87, 91 enhances z; increases duplication for recombination in w locus wch site wric spont Judd roo-in-copia 34, 136 eyes like wa; carries roo in copia insertion wRr EMS Pokholkova, Russian red 190 eyes vermilion, later 1976 almost wild type; ocelli colorless wrv#6 spont revertant 165 w+ revertant of w#6 wrv#12 spont 165 w+ revertant of w#12 wrvD5 118 revertant of wDZL ws10 spont Green, 1959 spontaneous 54 eyes white; not affected by su(wa); does not suppress z wsat spont Bridges, w331; satsuma 24-25, 54 eyes deep ruby; larval 33l26 Malpighian tubes nearly w+; does not suppress z wsey spont Erickson sepia-yellow 40-41 eyes lemon yellow with se at 25, white at 18 wsp1 Showell, spotted; sp-w 54, 57, facets yellowish to brown; 1944 89, 91 eyes with fine-grained 117, 120, 147, mottling; male darker than 187, 188 female; suppresses z; suppressed by su(wsp) wsp2 spont Mohler, 34, 48, eyes mottled like wsp; 56c22 129 wsp2/w and wsp/we uniform brown wsp3 X ray Green, 59a29 34, 52, 187 like wsp wsp4 spont LeFever sp-w4 34, 113, 147, 187 like wsp wsp55 17, 17a, 17b, 117, eye yellow; color lightened 187 by E(wa); variegated with mw; enhances z in XY male wsp81d EMS Green 34, 35 darker than wsp1; not suppressed by su(wsp) wsp+1 EMS Banerjee 9 revertant of wsp wsp+2 EMS Banerjee 9 revertant of wsp wsp+3 EMS Banerjee 9 revertant of wsp wsp+4 EMS Banerjee 9 revertant of wsp wspA / ray Alexandrov w76d 3 eyes darker and more speckled than wsp2; suppresses z wt spont Hyde, 14k2 tinged 77 eyes light pinkish; larval Malpighian tubes colorless wtache from Judd tache 79 like wzh, but red pigment wzm incomplete around circumference of eye wtuh spont Kuhn, tumorous head 103 like w1 Walker wu11 Fahmy, unstable 42, 48 eyes white; unstable Fahmy wudf spont Gethmann unstable deficient 48 like w1; wsp/wudf eyes pale lemon yellow; male viable wvC X ray Catcheside variegated of 74 eyes mottled; male In(1)3C1-2;20F + [in R(1)2] Catcheside viability zero -> fair; R(1)1A3-4;20F wvC/+ females may be N wvco Clausen variegated 111 eye mostly white with Tp(1;3)2B17-C1; cobbled red mottling 3C5-6; 77D3-5;81 wVD3 see wm258-21 wvr spont Mostashfi, venetian red 134 eyes white Kouliantz ww spont Kalisch, white 93-94 eyes white; mutates to In(1)3B2-C; (from 1980 wzm+, wzl, wzmz 4B4-C1 wzmz) wzmzz; may revert to normal X; does not suppress z wz from Judd, 1967 zeste 48, 88-89 white eyes; highly mutable wzm does not suppress z wzh from Judd, 1967 whalo 71, 79, 88, "halo" eyes with z in Tp(1;3)3C2-3; wzm 90, 114 homozygotes; mutable 3C6;61D wzl spont Becker, 1958 zl 12-13, enhances z; (from 63, 85, z wzl female has lemon wzm) 86 yellow eyes at 25-30; same female at 14 and male at 25-30 have yellow eyes with large red spots; z wzl male at 14 has red spots on yellow with salt-and-pepper mottling wzm spont Becker, 1959 zm 12-13, enhances z; eye color mottled (asym. 17, 17a, 63, with z and mw; exchange 85-86, 92, z wzm female like z wzl within w) 117, 147, 187 female at 25-30; same female at 14-17 has orange eyes with large red spots; z wzm male at 25-30 has yellow eyes with small red spots; at 14-17; male is w+; carries BEL wzm+ from Kalisch, 95 eyes of z wzm males like wzm Becker, z w+ at 14-17, wzmz 1970 duller at 25-30; eyes of wzmzz z wzm females between z w+ and z wzm wzmz spont Becker, 63, 94-95 eyes of z wzmz males (from 1959 red-yellow (pepper and wzm) salt mottling) at 14-17; males at 25-30 and all z females have lemon yellow eyes; mutates to wzm, wzm+, wzmzw, ww, wzmzz, wzmzrb wzmzrb from Kalisch, 95 at 28, z wzmzrb wzmz Becker, females have lemon 1970 yellow eyes, males have red to red brown eyes wzmzw spont Kalisch, 94-95 eyes of males and (from Becker, females white in genotypes wzmz) 1970 z wzmzw and z+ wzmzw; does not suppress z wzmzz spont Judd, 1967 63, 94-95 eyes of males with z (from deep yellow at 14-17, pale wzmz) yellow at 25-30; eyes of z females pale yellow at all temperatures wzs from Judd, 1967 88 eyes white; suppresses z wzm Table II synonym/ superscript allele origin discov. name ref ( comments cytology ______________________________________________________________________________________________________________________________________ *w13G2 X ray Gans eyes variegated T(1;2)3C3-5;56F *w48h mustard gas Lindsley, 122 like w1 see In(1)wm4 (from 48h13 wm4) *w48hS X ray Schultz, 48h w48h 31 like w1 *w51a spont Redfield, 51a 163 like w1 *w62d spont Mickey, 62d 127 like w1 (from wa) *w64 spont Whitten, 1964 31 like w1 *w258-43 X ray Demerec, 38k 31 male lethal *T(1;4)3C3-5;102F4-5 *w258-47 X ray Demerec, 39a 31 like w1 *w258-49 X ray Demerec, 39c 31 like w1 *w258-50 X ray Demerec, 39c 31 like w1 *w258-51 X ray Demerec, 39k 31 like w1 *w258-52 X ray Demerec, 40a 31 eyes white and rough *In(1)3C7-9;8E11-F1 (Sutton) *wa55k X ray Clark apr55k 28 like wa *wa57i spont Green, 57i11 waR57j 158-59 eye color between wa and (from wa) w+; more brown pigment; enhanced by su(f); does not suppress z *wa58l spont Green, 58l12 158-59 eye color between wa and w+; enhanced by su(f); does not suppress z *wa59k1 X ray Green, 59k1 158-59 eye color between wa and (from wa) w+; more brown pigment; enhanced by su(f), suppressed by su(wa); does not suppress z *wa59k9 X ray Green, 59k9 158-59 eye color between wa and (from wa) w+; more brown pigment; enhanced by su(f), suppressed by su(wa); does not suppress z *wa60a5 spont Sherwood, 158-59 eye color between wa and (from wa) 60a5 w+; more brown pigment; enhanced by su(wa); does not suppress z *wch2 cherry 54 like wch except no sex difference in eye color; does not suppress z *wch41j spont Ives, 41j9 78 like wch *wdil spont Ives, 33e31 w33e31; 156 eyes dilute red; dilute overlaps w+ *wdp2 chloroethyl Auerbach, deep purple 6, 51, like wdp; methane- 1957 54 does not suppress z sulfonate *wec spont Muller, 1918 ecru 28, 137 eyes very pale buff *wec2 54 like wec *wG X ray Goldschmidt Goldschmidt 49 like w1 see In(1)yG (in In(1)yG) *wm1 X ray Muller, 1927 mottled 50, 138 variegated for eye color *T(1;3) and N; XY male lethal, XYY male viable and sterile, with dark eye color *wm2 X ray Patterson, 138 eyes mottled; male sterile *T(1;3) 1929 *wm3 X ray Muller, 1929 138 only viable with normal X; *Dp(1;f)3C-D;19-20 w/Dp(1;f)wm3 -> mottling *wm11 X ray Panshin 151-53 eyes mottled *T(1;4)3C3-4;101A-D *wm52b12 X ray Ratty, 52b12 109 eyes variegated *Tp(1;2)1E5-F1;3C3-4; (in rst3) 20F;40-41 *wm52b13 X ray Ratty, 52b13 109 eyes variegated *Tp(1;4)2A2-3; (in rst3) 3C3-4;20F;101 *wm53a X ray Farnsworth, 109 eyes variegated *Tp(1;2)3B2-C1; 53a4 3C9-D1;40-41 *wm53e neutrons Mickey, 127 eyes variegated *T(1;2)3C3-4;20A2-3; 53e11 58F8-59A1 *wm53j X ray Bender, 53j 14 eyes mottled with small *In(1)1A;3C3-5; (in dark brown patches 20;20F;20F; In(1)EN) tentative *wm55b X ray Bender, 55b 14 eyes mottled with large *In(1)1A3-4;3C-5 [from pink or white patches 20;19F-20A1;20A1-F R(1)2] (inferred from origin) *wm258-31 X ray Demerec, 37l 31 eyes cream colored, *T(1;4)3C3-5; mottled; male viable 102F4-17 *wm258-32 X ray Demerec, 37l 31 eyes cream colored, *T(1;3)3C3-5;81 mottled; male viable *wm258-34 X ray Demerec, 38b 31 eyes cream colored *T(1;2)3C3-5;41A mottled; male viable *wm258-36 X ray Demerec, 38b 178 eyes cream colored *Tp(1;2)3C6-7;4C2-3; mottled; male viable 41A-B;41F5-6 *wm258-37 X ray Demerec, 38j 31 eyes mottled; male viable *T(1;2)3C3-4;40-41A *wm258-39 X ray Demerec, 38e 31 eyes cream colored, *T(1;2)3C3-5;40E-F mottled; male viable *wm258-40 X ray Demerec, 38e 31 eyes cream colored, *T(1;2)3C3-5;41 mottled, rough *wm258-44 X ray Demerec, 38k 31 eyes mottled; *T(1;2;3)3C3-4; male viable 4D2-E1; 56E1-F1;80D *wm258-53 X ray Demerec, 39l 178 eyes mottled; *T(1;4)3C1-2;101E-F male viable (101F-102F lost) *wm258-54 X ray Sutton, 40e 31 eyes cream colored, *T(1;3)3B2-C1; mottled; male lethal 19F2-20A1; 20E;63C7-8 *wmA X ray Stone 65-66 eyes variegated; male *T(1;4)3C2-3;101A2-3 (in viable and fertile YSX.YL) *wmCi X ray Cicak 150 eyes variegated with red and white facets *wmD1 X ray Dubinin 167 eyes variegated *T(1;2)3B;19-20;21F *wmDG1 Dubinin, 167 eyes variegated 3B inserted in Goldat chromocenter *wmDV4 Dubinin, 167 eyes mottled; male lethal; *T(1;4)3C3-7; Volotov wmD4/+ female N 3D;101A-D; (3C-3D missing) *wmMed Medvedev, 31 mottling on wa T(1;4) 1934 background *wM spont MacKendrick, of MacKendrick 124, eye color almost w+ (less (from wa) 1955 158-59 red pigment); wM/w like wco (a bit darker); viable and fertile; enhanced by su(wa) and su(f); does not suppress z *wM59 spont Muller, 59d of Muller 128 eyes cream colored, darkening a bit with age *wp spont Steinberg, pearl 24, 175 eyes very pale (lighter than 37b17 wt); larval Malpighian tubes colorless *ws1 spont Green, 1959 spontaneous 54 eyes white; does not suppress z *ws2 spont Green, 1959 54 eyes white; does not suppress z *ws3 spont Green, 1959 54 eyes white; does not suppress z *ws4 spont Green, 1959 54 eyes white; does not suppress z *ws5 spont Green, 1959 54 eyes white; does not suppress z *ws9 spont Green, 1959 54 eyes white; does not suppress z *wvD1 X ray Demerec, variegated 31 eyes variegated red *Dp(1;4)3C1-4; 33j19 of Demerec and white; male fertile 101A-D *wvD2 X ray Demerec, 31 eyes with fine-grained *T(1;2;4)3C4-5;18F; 33k27 variegation of cream 38;101A-C with dark spots; female occasionally variegated for rst *wvD4 X ray Demerec, 31 wvD4/+ female mottled; *T(1;2)3D6-E1;40F 33k2 XY, wvD4 male mottled, usually lethal; XYY male sterile *wX1 X ray Green, 1959 from X 54 eyes white; irradiation does not suppress z *wX2 X ray Green, 1959 54 eyes white; does not suppress z *wX3 X ray Green, 1959 54 eyes white; does not suppress z *wX4 X ray Green, 1959 54 eyes white; does not suppress z *wX5 X ray Green, 1959 54 eyes white; does not suppress z *wX6 X ray Green, 1959 54 eyes white; does not suppress z *wX8 X ray Green, 1959 54 eyes white; does not suppress z *wX16 X ray Green, 1959 54 eyes white; suppresses z ( 1 = Alexandrov, 1971, DIS 46: 71, 72; 2 = Alexandrov, 1972, DIS 48: 88, 133; 3 = Alexandrov, 1982, DIS 58: 7-8, 9-10, 10-12; 4 = Alexandrov and Soluyanova, 1974, DIS 51: 32; 5 = Anxolabehere and Periquet, 1973, DIS 50: 21; 6 = Auer- bach, 1957, DIS 31: 107-09; 7 = Baker, 1963, Am. Zool. 3: 57-69; 8 = Baker and Spofford, 1959, Univ. Texas Publ. 5914: 135-54; 9 = Banerje, Hazra, and Sen, 1978, Mutat. Res. 50: 309-15; 10 = Beadle and Ephrussi, 1936, Genetics 21: 230; 11 = Beckendorf, 1983, Nucleic Acids Res. 11: 737-51; 12 = Becker, 1959, DIS 33: 82; 13 = Becker, 1960, Genetics 45: 519-34; 14 = Bender, 1955, DIS 29: 69; 15 = Bingham, 1980, Genetics 95: 341-53; 15a = Bingham and Chapman, 1986, EMBO J. 5: 3343-51; 16 = Bingham and Zachar, 1985, Cell 40: 819-25; 17 = Birchler, 1986, Genetics 113: s47; 17a = Birchler and Hiebert, 1989, Genetics 122: 129-38; 17b = Birchler, Hiebert, and Rabinow, 1989, Genes Dev. 3: 73-84; 18 = Bolen, 1931, Am. Nat. 65: 417- 22; 19 = Bowman, 1967, DIS 42: 73; 20 = Bowman, 1968, DIS 43: 91; 21 = Bowman, 1969, Mutat. Res. 7: 409-15; 22 = Bowman and Green, 1964, Genetics 50: 237; 23 = Braver, 1953, DIS 27: 86; 24 = Brehme and Demerec, 1942, Growth 6: 351-56; 25 = Bridges, 1935, DIS 3: 18; 25a = Bridges, 1938, DIS 9: 114; 25b = Bucheton, Paro, Sang, Pelisson, and Finnegan, 1984, Cell 38: 153-63; 26 = Buzzati-Traverso, 1943, Rend. Ist. Lombardo Sci. Lettere, Pt. I: Class. Sci., Mat. e Nat. 77: 61-64; 27 = Carbonara and Gehring, 1985, Mol. Gen. Genet. 199: 1-6; 28 = Clark, 1956, DIS 30: 71; 29 = Collins and Rubin, 1982, Cell 30: 71-79; 29a = Collins and Rubin, 1983, Nature (London) 303: 259-60; 30 = Collins and Rubin, 1984, Nature (London) 308: 323-27; 31 = CP627; 32 = Craymer, 1980, DIS 55: 197-200; 33 = Curry, 1938, DIS 9: 114; 34 = Davis, Shen, and Judd, 1987, Proc. Nat. Acad. Sci. USA 84: 174-78; 35 = Davison, Chapman, Wedeen, and Bingham, 1985, Genetics 110: 479-94; 36 = Demerec, 1940, Genetics 25: 618-27; 37 = Demerec and Slizynska, 1937, Genetics 22: 641-49; 38 = Dunn, 1935, Hereditas 21: 113- 18; 39 = Ephrussi and Herold, 1945, Genetics 30: 62-70; 40 = Erickson, 1974, DIS 51: 22; 41 = Erickson, 1983, DIS 59: 146; 42 = Fahmy and Fahmy, 1983, Cancer Res. 43: 801- 07; 43 = Fahmy and Fahmy, 1984, Carcinogesis and Mutagenesis, 4: 437-47; 44 = Farmer and Fairbanks, 1984, Genetics 107: s30; 45 = Farmer and Fairbanks, 1986, DIS 63: 50-51; 46 = Gans, 1953, Bull. Biol. France Belg., Suppl. 38: 1-90; 47 = Gersh, 1967, Genetics 56: 309-19; 48 = Gethmann, 1971, Mol. Gen. Genet. 114: 144-55; 49 = Goldschmidt, 1945, Univ. Calif. Publ. Zool. 49: 522; 50 = Gowen and Gay, 1933, Proc. Nat. Acad. Sci. USA 19: 122-26; 51 = Green, 1958, DIS 32: 88; 52 = Green, 1959, DIS 33: 94; 53 = Green, 1959, Genetics 44: 1243-56; 54 = Green, 1959, Heredity 13: 303-15; 55 = Green, 1959, Nature (London) 184: 294; 56 = Green, 1959, Proc. Nat. Acad. Sci. USA 45: 549-53; 57 = Green, 1959, Z. Indukt. Abstamm. Vererbungsl. 90: 375-84; 58 = Green, 1960, Genet. Res. 1: 452-61; 59 = Green, 1967, Genetics 56: 467-82; 60 = Green, 1968, Proc. Int. Congr. Genet., 12th, Vol. 1: 88; 61 = Green, 1969, Genetics 61: 423-28, 429-41; 62 = Green, 1973, Genetics 73 (supplement): 187-94; 63 = Green, 1976, The Genetics and Biology of Drosophila (Ashburner and Novitski, eds.). Academic Press, London, New York, San Francisco, Vol. 1b, pp. 929-46; 64 = Griffen and Stone, 1938, Genetics 23: 149; 65 = Griffen and Stone, 1939, Genetics 24: 73; 66 = Griffen and Stone, 1940, Texas Univ. Publ. 4032: 201-07; 66a = Grimwade, Muskavitch, Welshons, Yedvobnick and Artavanis-Tsakonis, 1985, Dev. Biol. 107: 503-19; 67 = Hadorn and Mitchell, 1951, Proc. Nat. Acad. Sci. USA 37: 650-65; 68 = Hanly, 1963, DIS 38: 30; 69 = Hartl and Green, 1970, Genetics 65: 449-56; 70 = Hayman and Maddern, 1969, DIS 44: 50; 71 = Hazelrigg, Levis, and Rubin, 1984, Cell 36: 469-81; 72 = Hess, 1979, pers. comm.; 73 = Hessler, 1961, Genetics 46: 463-84; 74 = Hinton, 1955, Genetics 40: 952-61; 75 = Hollander, 1960, DIS 34: 50; 76 = Hollander, 1962, DIS 36: 78; 77 = Hyde, 1916, Genetics 1: 535-80; 78 = Ives, 1942, DIS 16: 58; 79 = Jack and Judd, 1979, Proc. Nat. Acad. Sci. USA 76: 1368-72; 80 = Judd, 1955, Genetics 40: 739-44; 81 = Judd, 1957, Genetics 42: 379-80; 82 = Judd, 1958, Proc. Intern. Congr. Genet., 10th, Vol. 2: 137; 83 = Judd, 1959, Genetics 44: 34-42; 84 = Judd, 1961, Proc. Nat. Acad. Sci. USA 47: 545-50; 85 = Judd, 1963, Proc. Intern. Congr. Genet. 11th, Vol. 1: 3-4; 86 = Judd, 1964, DIS 39: 59-60; 87 = Judd, 1964, Genetics 49: 253-65; 88 = Judd, 1967, Genetics 56: 569; 89 = Judd, 1969, Genetics 61: s29; 90 = Judd, 1975, The Eukaryotic Chromosome (Peacock and Brock, eds.). Aust. Nat. Univ. Press, Canberra, pp. 169-84; 91 = Judd, 1976, The Genetics and Biology of Drosophila (Ashburner and Novitski, eds.). Academic Press, London, New York, San Francisco, Vol. 1b, pp. 767-99; 92 = Judd, 1987, Structure and Function of Eukaryotic Chromosomes (Hennig, ed.). Springer-Verlag, Berlin, Heidelberg, pp. 81-94; 93 = Kalisch, 1970, Mol. Gen. Genet. 107: 335-50; 94 = Kal- isch, 1980, DIS 55: 206-07; 95 = Kalisch and Becker, 1970, Mol. Gen. Genet. 107: 321-35; 335-50; 96 = Kalmus, 1943, J. Genet. 45: 206-13; 97 = Karess and Rubin, 1982, Cell 30: 63-69; 98 = Kaufman, 1969, DIS 44: 44; 99 = Kaufman, 1970, DIS 45: 34; 100 = Kaufman, Tasaka, and Suzuki, 1973, Genetics 75: 299-321; 101 = Kennison and Ripoll, 1981, Genetics 98: 91-103; 102 = Kidd, 1966, DIS 41: 60; 103 = Kuhn and Walker, 1980, DIS 55: 207; 104 = Lancefield, 1918, Am. Nat. 52: 264-69; 105 = Lee, 1973, Aust J. Biol. Sci. 26: 903-09; 106 = Lefevre, 1949, DIS 23: 59; 107 = Lefevre, 1951, DIS 25: 71; 108 = Lefevre, 1952, DIS 26: 66; 109 = Lefevre, 1953, DIS 27: 57; 110 = Lefevre, 1963, DIS 37: 49-50; 111 = Lefevre, 1981, Genetics 99: 461-80; 112 = Lefevre and Wilkins, 1966, Genetics 53: 175-87; 113 = LeFever, 1973, DIS 50: 109-11; 114 = Levis, Bingham, and Rubin, 1982, Proc. Nat. Acad. Sci. USA 79: 564-68; 115 = Levis, Collins, and Rubin, 1982, Cell 30: 551-65; 116 = Levis, Hazelrigg, and Rubin, 1985, Sci- ence 229: 558-61; 117 = Levis, O'Hare, and Rubin, 1984, Cell 38: 471-81; 118 = Levis and Rubin, 1982, Cell 30: 543-50; 119 = Lewis, 1952, Proc. Nat. Acad. Sci. USA 38: 953-61; 120 = Lewis, 1956, Genetics 41: 651; 121 = Lewis, 1959, Genetics 44: 522; 122 = Lindsley, 1949, DIS 23: 60; 123 = MacKendrick, 1953, DIS 27: 100; 124 = MacKendrick, 1958, DIS 32: 82; 125 = MacKendrick and Pontecorvo, 1952, Experientia 8: 390-91; 126 = Mainx, 1938, Z. Indukt. Abstamm. Vererbungsl. 75: 256-76; 127 = Mickey, 1963, DIS 38: 29; 128 = Mischaikow, 1959, DIS 33: 98; 129 = Mohler, 1956, DIS 30: 78-79; 130 = Morgan, 1910, Sci- ence 32: 120-22; 131 = Morgan and Bridges, 1916, Carnegie Inst. Washington Publ. No. 237: 25, 28, 51; 132 = Morgan, Bridges, and Sturtevant, 1925, Bibliog. Genet. 2: 218; 133 = Mossige, 1953, DIS 27: 59; 134 = Mostashfi and Kouli- antz, 1969, DIS 44: 51; 135 = Mount, Green, and Rubin, 1988, Genetics 118: 221-34; 136 = Mount and Rubin, 1985, Mol. Cell Biol. 5: 1630-38; 137 = Muller, 1920, J. Exp. Zool. 31: 433-73; 138 = Muller, 1930, J. Genet. 22: 299- 334; 139 = Muller, 1932, Proc. Intern. Congr. Genet., 6th, Vol. 1: 234; 140 = Muller, 1935, J. Genet. 30: 407-14; 141 = Muller, 1944, DIS 18: 57; 142 = Muller, 1946, DIS 20: 68; 143 = Najera, 1984, DIS 60: 241-42; 144 = Neeley, 1971, DIS 46: 43; 145 = Nicoletti, 1960, DIS 34: 52-53; 146 = Nolte, 1954, DIS 28: 77; 147 = O'Hare, Murphy, Levis, and Rubin, 1984, J. Mol. Biol. 180: 437-55; 148 = Oliver, 1935, DIS 3: 28; DIS 4: 12; 149 = Osgood and Lacy, 1985, Genetics 110: s36; 150 = Oster, 1957, DIS 31: 150; 151 = Panshin, 1938, Nature (London) 142: 837; 152 = Panshin, 1941, DIS 15: 33-34; 153 = Panshin and Khvostova, 1938, Biol. Zh. (Moscow) 7: 359-80; 153a = Pastink, Schalet, Vreeken, Paradi, and Eeken, 1987, Mutat. Res. 177: 101-15; 153b = Pastink, Vreeken, Schalet, and Eeken, 1988, Mutat. Res. 207: 23-28; 153c = Pastink, Vreeken, and Vogel, 1988, Mutat. Res. 199: 47-53; 154 = Pelisson, 1981, Mol. Gen. Genet. 183: 123-29; 155 = Phillips and Forrest, 1980, The Genetics and Biology of Drosophila (Ashburner and Wright, eds.). Academic Press, London, New York, San Francisco, Vol. 2d, pp. 541-623; 156 = Plough and Ives, 1934, DIS 1: 31; 157 = Rasmuson, 1962, Hereditas 48: 587-611; 158 = Rasmuson, Green, and Ewertson, 1960, Hereditas 46: 635-50; 159 = Rasmuson and Rasmuson, 1961, Hereditas 47: 619-30; 160 = Ratty, 1954, Genetics 39: 513-28; 161 = Rayle, 1968, DIS 43: 62; 162 = Rayle and Green, 1969, Genetica 39: 497-507; 163 = Redfield, 1952, DIS 26: 68; 164 = Reuter and Wolff, 1981, Mol. Gen. Genet. 182: 516-19; 165 = Rubin, Kidwell, and Bingham, 1982, Cell 29: 987-94; 166 = Ryo, Yoo, Fujikawa, and Kondo, 1985, Genetics 110: 441-51; 167 = Sacharov, 1936, Biol. Zh. (Moscow) 5: 293-302; 168 = Safir, 1913, Biol. Bull. 25: 45-51; 169 = Safir, 1916, Genetics 1: 584-90; 170 = Sang, Pelisson, Bucheton, and Finnegan, 1984, EMBO J. 3: 3079-85; 171 = Simmons and Lim, 1980, Proc. Nat. Acad. Sci. USA 77: 6042-46; 172 = Spofford, 1958, Proc. Intern. Congr. Genet. 10th, Vol. 2: 270; 173 = Spofford, 1959, Proc. Nat. Acad. Sci. USA 45: 1003-07; 174 = Spofford, 1961, Genetics 46: 1151-67; 175 = Steinberg, 1937, DIS 8: 11; 176 = Stern, 1969, Genet- ics 62: 573-81; 177 = Sturtevant and Beadle, 1939, An Introduction to Genetics, W.B. Saunders Co., Philadelphia, p. 64 (fig.); 178 = Sutton, 1940, Genetics 25: 534-40, 628-35; 179 = Tartof, Hobbs, and Jones, 1984, Cell 37: 869-78; 180 = Timofeeff-Ressovsky, 1932, Biol. Zen- tralbl. 52: 468-76; 181 = Valade del Rio, 1982, DIS 58: 144-45; 182 = Valencia, 1966, DIS 41: 58; 183 = Welshons and Keppy, 1981, Mol. Gen. Genet. 181: 319- 24; 184 = Welshons and Nicoletti, 1963, DIS 38: 80; 185 = Whitney and Lucchesi, 1972, DIS 49: 35; 186 = Wright and Hanley, 1966, Science 152: 533-35; 187 = Zachar and Bingham, 1982, Cell 30: 529-41; 188 = Zachar, Chapman, and Bingham, 1985, Cold Spring Harbor Symp. Quant. Biol. 50: 337-46; 189 = Zachar, Davison, Garza, and Bingham, 1985, Genetics 111: 495-515; 190 = Zhimulev, Belyaeva, Khu- dyakov, and Pokholkova, 1980, DIS 55: 211. | Heteroalleles with wsp have brown eyes at eclosion, comple- menting wsp; eyes somewhat darker after 4-5 days. / Heteroalleles with wsp have yellow-orange, orange, or orange-brown eyes at eclosion; mottling only on first day. ` Heteroalleles with wsp usually have orange-brown eyes at eclosion; mottling almost gone after first day; eyes slowly darken. - Heteroalleles with wsp have yellow eyes at eclosion, usually with mottling; eyes become orange after 4-5 days, but not brown. cytology: Placed in 3C2 by Schultz and also by Lefevre and Wil- kins (1966, Genetics 53: 175-987) on the basis of rearrange- ment breakpoints [In(1)w4 = In(1)3C1-2;20F, for example]. An electron microscope study of In(1)z+64b9 = In(1)3C1-2;12B9-10 shows a faint band to the right of 3C1 consisting of the left- most border of 3C2 plus material from 12B9 [Sorsa, Green, and Beerman, 1973, Nature (London) New Biol. 245: 34-37]; this band is not seen in most preparations of normal chromosomes. Judd (1976, 1987) suggests that w+ is located in a very small band between 3C1 and 3C2. Goldberg et al. (1982) showed that the 3C1-2 break of In(1)z+64b9 is proximal to the white locus on the molecular map (about 10 kb to the right of the wa insertion of copia). molecular biology: The wa allele was the first gene cloned by "transposon tagging", using previously cloned transposing ele- ment copia (Bingham et al., 1981). The wa clone was extended on both sides by chromosome walking and a segment of 14 kb was found to contain all of the white locus sequences essential to the w+ eye-color phenotype (Levis et al., 1982). The white locus was also cloned by Goldberg et al. (1982), using a large TE containing wa and rst inserted at 87A7 next to previously cloned sequences (genes encoding Hsp70). Pirrotta et al. (1983) later cloned white by microdissection of the 3C2 region of the salivary chromosomes. A physical map of the white locus was prepared after cleav- ing DNA from various mutants with restriction endonucleases (Levis et al., 1982; Goldberg et al., 1982); transcripts were identified, the major one being a 2.6 kb poly-(A)+ RNA found in embryos, larvae, pupae, and adults (O'Hare et al., 1983; Pirrotta et al., 1983; Fjose et al., 1984; Pirrotta and Brockl, 1984). The nucleotide sequence of more than 14 kb of white DNA and the probable structures of the introns and exons were determined (O'Hare et al., 1984). The presence of at least four introns is indicated (Pirrotta and Brockl, 1984); a 3 kb intron separates the proximal (centromere) sequences from the distal sequences (O'Hare et al., 1984). DNA sequences required for normal expression of w+ were found to be within a 9.9 kb segment of the gene, with regulatory elements in the 5' flanking region (Levis et al., 1985; Pirrotta et al., 1985); these regions include sequences for Malpighian tubule expres- sion and testis sheath pigment as well as those regulating the pattern of eye pigmentation and zeste interaction. Insertions associated with w mutations were found throughout most of the region of the 2.6 kb w+ transcript as well as 6 kb upstream. Alleles associated with these insertions or with molecular deletions or duplications are listed on the figure and table on the following page. The mottler of white locus interacts with white alleles that are transposon-insertion mutants, producing a variegated eye color (Birchler et al., 1989). All of the w sites involved are within the structural part of the gene. w+ (red-eyed) flies were produced by P-element-mediated germ-line transformation, a 12-14 kb segment spanning the w+ locus (in a P-transposon vector) being inserted in various new chromosomal locations (Gehring et al., 1984; Hazelrigg et al., 1984) and with various amounts of upstream sequences (Levis et al., 1984; Pirrotta et al., 1985). These w+ transformants show the same tissue-specific transcript accumulation as wild-type flies. They also show dosage compensation, but differ markedly in their interactions with z. Temperature shocks given at various times in development to white transformants under the control of heat shock regulatory sequences indicate the period of maximum expression of white to be in the first two days of the pupal period (Steller and Pirrotta, 1985). Extensive amino acid similarity has been found between the putative protein products of the white and brown genes, both involved in the pteridine pathway (Dreesen et al., 1988) and the protein products of the white and scarlet genes, both involved in the ommochrome pathway (Tearle et al., 1989). While the amount of detectable eye pigment variation among 64 X chromosome lines from three collections in North Caro- lina, Texas, and Fukuoka, Japan, was not large, these lines showed a very large amount of molecular variation in a 45-kb region of the white locus, a total of 109 polymorphisms being found (Miyashita and Langley, 1988). other information: The original TE35A [= TE146(Z)] which car- ries two copies of w+rst+ in tandem order (w+rst+w+rst+) is prevented by certain rearrangements on the homologous chromo- some from interacting with z1, but a derivative of TE35A which carries two copies of w+rst+ in the reversed order (rst+w+w+rst+) is not affected by these rearrangements in regard to its interaction with z1 (Gubb et al., 1990). white molecular map Modified from data supplied by O'Hare, Murphy, Levis, and Rubin, 1984, J. Mol. Biol. 184: 437-55. By D. Conner. . allele mut. agent size of agent map site (kb) ( ref | ____________________________________________________________________________ w1 Doc 5 kb +3.71 14, 16 w5a translocation break +4.38 to +4.44 12 w13D1 deletion -3.0 to -0.7 12 w17D2 deletion -3.0 to -0.7 12 w25B2 deletion -1.5 to -0.7 12 w31D1 translocation break -3.0 to -0.7 12 w35A2 deletion -3.0 to -0.7 12 w41C1 deletion -3.0 to -0.7 12 w53A dup (3A-3C) +2.0 to +4.8 12 + deletion w66A deletion 12 w73A1 deletion -3.0 to -1.5 12 w83B translocation break -3.0 to -0.7 12 w108 deletion +3.2 to +4.4 13 w123D inversion break (3C) -0.7 to +0.4 12 in Tp(3;1) w132 insertion 10 kb +0.4 in large 13 intron w#6 P 1.1 kb -2.02 11, 14 w#12 P 1.6 kb -0.51 11, 14 wa / copia insertion 5.0 kb 0 2, 7, 8, 10, in intron 2 11, 16 wa4 / BEL insertion 7.3 kb 8, 16 in intron 2 waM / insertion in 2.3 kb 10 copia 5' LTR waR84h / insertion in 10 copia 3' LTR wbf / roo 8.7 kb -1.13 8, 11, 16 wbl blood 6 kb -0.01 1 wc FB in wi 9 kb +0.19 3, 11 wch pogo in Doc 0.2 kb +3.71 O'Hare, 16 wDZL FB 13 kb +9.77 9, 11, 16 we pogo in Doc 0.2 kb +3.71 8 we2 pogo in Doc 0.2 kb +3.71 O'Hare we59 F +3.76 O'Hare wh / roo in Doc 5.7 kb +3.71 11 whd80k P -2.03 8, 11, 14 whd81b11 copia -1.3 11, 14 wi dup of 2.96 kb -0.17 to +2.80 3, 11 intron 1 wi+A F 3 kb +0.8 8, 11 wIR1 I 5.4 kb -1.45 6, 15 wIR2 I into Doc 5.4 kb 6, 15 wIR3 I 5.4 kb -1.45 6, 15 wIR4 I 5.4 kb -1.45 6, 15 wIR5 I 5.4 kb -2.2 6, 15 wIR6 I 5.4 kb +3.2 6, 15 wIR7 deletion -1.17 to 6, 15 outside w wIR8 deletion -1.29 to -3.45 6, 15 wric roo in copia 4 wsp1 ` roo 8.7 kb +4.92 8, 11, 16 wsp2 deletion in 16 regulatory region wsp3 deletion +5 to +22 16 wsp4 deletion in +4.78 to +5.85 11, 16 regulatory region wsp55 /- retrotransposon 5.8 kb +3 to +5 8, 16 insertion wsp81d5 deletion 5 wzm BEL 6.0 kb +3.43 8, 11, 16 ( Origin = insertion of wa copia; "-" values to left (telomere) end; "+" values to right (centromere) end. | 1 = Bingham and Chapman, 1986, EMBO J. 5: 3343-51; 2 = Bingham and Judd, 1981, Cell 25: 705-11; 3 = Collins and Rubin, 1982, Cell 30: 71-79; 4 = Davis, Shen, and Judd, 1987, Proc. Nat. Acad. Sci. USA 84: 174-78; 5 = Davison, Chapman, Wedeen, and Bingham, 1985, Genetics 110: 479-94; 6 = Fawcett, Lister, Kellett, and Finnegan, 1986, Cell 47: 1007-15; 7 = Gehring and Paro, 1980, Cell 19: 897-904; 8 = Levis, O'Hare, and Rubin, 1984, Cell 38: 471-81; 9 = Levis and Rubin, 1982, Cell 30: 543-50; 10 = Mount, Green, and Rubin, 1988, Genetics 118: 221-34; 11 = O'Hare, Murphy, Levis, and Rubin, 1984, J. Mol. Biol. 180: 437-55; 12 = Pastink, Schalet, Vreeken, Paradi, and Eeken, 1987, Mutat. Res. 177: 101-15; 13 = Pastink, Vreeken, and Vogel, 1988, Mutat. Res. 199: 47-53; 14 = Rubin, Kidwell, and Bingham, 1982, Cell 29: 987-94; 15 = Sang, Pelisson, Buche- ton, and Finnegan, 1984, EMBO J. 3: 3079-85; 16 = Zachar and Bingham, 1982, Cell 30: 529-41. Also see references in Table 1 of Judd, 1987, Structure and Function of Eukaryotic Chromosomes (Hennig, ed.). Springer-Verlag, Berlin, Heidel- berg, pp. 81-94 and Birchler, Hiebert, and Rabinow, 1989, Genes Dev. 3: 73-84. / Shows mw mottling (Birchler et al., 1989). wa suppressed by Doa (Rabinow and Birchler, 1989, EMBO J. 8: 879-89). ` In spite of the variegated pattern of spots in the eye in this mutant, the transcript is like that of w+ in size, structure, and amount (Levis et al., 1984; Pirrotta and Brockl, 1984, EMBO J. 3: 563-68). - w+ mRNA not detected in this allele. wsp55 enhanced by Doa (Rabinow and Birchler, 1989).