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A Third Sex-Linked Lethal Factor in Drosophila T

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A Third Sex-Linked Lethal Factor in Drosophila T A THIRD SEX-LINKED LETHAL FACTOR IN DROSOPHILA T. H. MORGAN Department of ZoBlogy, Columbia University THREE FIGURES In an experiment (reported in the Jour. Exp. Zool., page 390, November, 1911) in which three sex-linked characters were in- volved, namely, white eyes, vermilion eyes and miniature wings, certain results were obtained involving double crossing-over. The more recent results indicate that double crossing-over was excessive in this experiment.l A white miniature male had been mated to a vermilion pink (‘orange’) female. The daughters were wild type (red, long) and the sons vermilion (long). The two sex- chromosomes of these daughters contain respectively the genes for white (w), red (V), and miniature (m); and for red (W), and vermilion (v), and long (M). All later work has shown the order of the genes here involved is white, vermilion, miniature, as shown in diagram 1. On this basis, the results of the previous experiment in terms of cross-overs would be as follows: Total Per cent Non-cross-overs (vermilion and ver. pink males) 333 74.7 Single cross-overs between white and vermilion (min. and min. pink males). 81 18.2 Single cross-overs between vermilion and min. (verm. min. and ver. min. pink males). 11 2.4 Double cross-overs (wild type and pink males). 21 4.7 Instead of giving fewer double cross-overs than singles, accord- ing to expectation, the number of doubles was twice that of singles, and for numbers as large as these this difference cannot be due to 1 In the analysis on page 392 the large C in ‘sperm no X’ should be small c. This change adds eight classes to the white males and eliminates the same number from the colored classes. The expected numbers in the male classes at the top of page 393 should read downwards: 3 - 3 - 1 - 1 - 8 - 3 - 3 - 1- 1-8. 315 THE JOURNAL.OF EXPERIMENTAL ZO~LOGY,VOL. 17, NO. 3 OCTOBER. 1914 316 T. H. MORGAN Diagram 1 chance. This experiment has been repeated, therefore, in small mass cultures and has given, in Fz,the totals shown in table 1. TABLE 1 Females WILD TYPE I VERMILION I PINK I VERY. PISK 1428 1465 469 548 VIURUILION 2:;. MlN. p"I'$ ':&M' 'zz':iii PINK WEITE 501 161 251 84 14 35 7 0 314 1166 The females conform to expectation. The males show many irregularities that will be discussed in another connection. Classi- fied as in the original experiment, the males give : Totals Per cent Non-cross-overs ................................ 662 63.8 Single cross-overs between white and vermilion. 335 31.8 Single cross-overs between verm. and minia- ture ....................................... 49 4.7 Double cross-overs ............................. 7 0.7 The totals of these new results for the males approach very closely to expectation both for the amount of double crossing-over and also for the distances apart of the genes, despite other inequali- ties that appear in the results. In the experiment as actually carried out a non-sex-linked gene, namely, the gene for pink eyes, is involved. The additional classes that are thus introduced are ignored in the preceding calcu- lations in which pink and not-pink classes are combined. If we take pink into consideration the following relations will be found. The gene for pink lies in another chromosome, and since the F1 A THIRD SEX-LINKED LETHAL 317 female is heterozygous for pink every egg after the polar bodies are extruded will contain either the pink factor or its normal allelomorph. The male has two classesof sperm, viz., one with and the other without the pink factor. In all cases except white (which remains white in external appearance in either case) the combination gives two classes in the ratio of 3:l. The relation of pink to not-pink are everywhere normal except in the following case. The single cross-overs between vermilion and miniature are vermilion miniature (14) and vermilion miniature pink (35). Instead of the expected 3 :1 ratio the relation is almost reversed. I can offer no explanation of the relative deficiency in the vermilion miniature class or the relative excess of the vermilion miniature pink. We may assume that the double cross-over class included in white miniature is very small and negligible; if so, then the non- cross-over class white miniature should equal the sum of the ver- milion and vermilion pink, which is 662. In fact, it is nearly double (1166) that number. This disturbance, as will now be shown, is due to a lethal factor present in the stock. THE LOCATION OF LETHAL I11 In the experiment, made as explained above to test double crossing-over, only about half as many males as females appeared in some of the cultures. This kind of rmult we have come to associate with a lethal factor, and further tests madewith some of the Fz females show, in fact, the presence of a new lethal. Table 2 gives individually the ten original mass cultures (p. 318). Since the cultures were made in mass they can not be used to study lethal factors, for, the offspring may have arisen from both lethal-bearing and non-lethal-bearing females. The high sex- ratio was not noticed until the experiment was nearly finished, and then it was possible to obtain only a few females.2 In three cases 2 We have known that non-disjunction occurs in several of our stocks, notably in the white. It occurred in the mother in the experiment just described, for when vermilion pink females were mated to white miniature males there were produced: Venn. '2 Red 0 Venn.$ White C? 2 82 46 2 The two vermilion females and the two white males must have arisen through 318 T. H. MORGAN TABLE 2 Females VERMILION WILD TYPE VERMILION PINK PIMK I. ........................ 1 28 123 39 51 11......................... 151 1.54 37 58 111. ......................... 173 169 6l 76 IV. ........................ 86 89 37 36 v ......................... 250 23 1 51 81 VI......................... 84 84 41 44 VII. ........................ 90 93 32 33 VIII. ......................... 94 96 28 31 IX ......................... 189 228 68 58 x ......................... 184 188 75 80 1428 1465 469 548 Males I- MIR. IERM. VHITE MlN. 'INK VHITE PINK YIN. MIX. - - - - - - I .......... 24 9 3 33 120 11.. ......... 30 5 2 36 142 111.. ......... 29 9 1 29 133 IV. .......... 10 2 1 16 37 v ........... 103 46 17 2 54 193 VI........... 12 11 7 2 16 78 VII. .......... 12 5 13 66 VIII. .......... 35 17 10 1 17 74 IX. ......... 28 10 2 45 173 x ........... 44 10 2 41 55 150 501 1161 I5 1 84 14 35 I 1 314 1166 at least, viz., IV, VIII and X, the sex-ratios were again obtained from Fzfemales. Further tests were made of the stock derived from the F3 lethal females in order to locate the lethal or lethals present. non-disjunction. One of the vermilion females was tested by breeding to a pink male. She gave: Red 0 Pink 0 Verm .c7 Verm. pink3 Red3 66 61 43 19 2 The two red males arc again due to non-disjunction. A THIRD SEX-LINKED LETHAL 319 Vermilion females were out-crossed' to club males. From those sets that gave a sex-ratio, daughters (red eyed) were mated singly to their vermilion-eyed brothers. About half of the females gave no sex-ratios, as follows: All females Club8 Verm.3 Clubverm.8 Wild typed 687 219 314 58 64 The cultures that gave sex-ratios follow (table 3) : ALL FEMALES CLUB 8 WILD TYPE # CLUB VGEM. 8 VEBM. # 146 53 3 1 176 59 7 3 145 34 10 2 1 121 36 8 5 588 182 28 11 1 The smallest class of males is vermilion, which we assume there- fore to be the double cross-over class. Since the lethal went in with the vermilion female one of the chromosomes of her daughter will contain the genes for lethal and for vermilion; and the homol- ogous sex chromosome will contain the gene for club. The rela- tion of the two chromosomes with the normal allelomorphs of these genes are shown in diagram 2. c1 &3 V X -I Cl 13 V Diagram 2 In order that vermilion become the double cross-over class the lethal must be between club and vermilion, as in the diagram. The chromosome distances calculated from the results are as follows: Club lethal 111.. ................................................. 13.0 Lethal 111 vermilion ............................................... 5.4 Club vermilion .................................................... 18.0 330 T. H. MORGAN Other vermilion females were out-crossed to eosin miniature males. From the cultures that showed sex-ratios females were back-crossed singly to eosin miniature males. The back-cross cultures that gave sex-ratios are seen in table 4. The results of the experiment confirm the location of the genes in diagram 1, as the following chromosomal distances show:, Eosin lethal 111.. ................................... Eosin vermilion.. ................................... Eosin miniature.......... ... .......... Lethal I11 vermilion., ............................................ 7 .O Lethal 111 miniature.. ............................................ 9.3 Vermilion miniature. ............................................ 3.3 TABLE 4 Females EOSlW SIN. I WILD TmE I EOSIN I MINIATURE 78 104 33 24 90 89 36 33 82 84 37 45 101 77 31 30 105 97 31 43 127 86 35 33 62 62 29 13 60 53 36 38 58 51 24 30 I 763 703 292 I 289 Males A THIRD SEX-LINKED LETHAL 321 The small numbers in the first experiment gave a distance between lethal I11 and vermilion of 5.4; the second gave 7. The data of both experiments gave 6.8.
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