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UBC Theses and Dissertations

A study of a possible new mutation, synpalpi, occurring in drosophila melanogaster Cole, Kathleen Margaret 1947

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THE STUDY OP A POSSIBLE HEW MUTATION, SYNPALPI, OCCURRING IN DROSOPHILA MELANOGASTER. BY K A J f f W a r MARGARET COLE A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF THE UNIVERSITY OF BRITISH COLUMBIA APRIL, 1947. MASTER OF ARTS IN THE DEPARTMENT OF BIOIOGY AND BOTANY ACZHOTSLEDGaiEin? S The writer wishes to express her sincere appreciation to Mrs. Ruth Fields Brink. Assistant Professor, for al l the assistance given during the following experimental study. Mrs. Brink* who dis-covered the mutation under investigation, was always ready to advise and showed keen interest and kindness at al l times* Gratitude is extended to Br* A. H. Hutchinson* - Head of the Department* who made available a l l necessary materials and equipment and who was most encouraging and helpful. The many useful suggestions given by Dr. V. C. Brink* Associate Professor* Department of Agronomy* are gratefully acknowledged. TABLE OF PORTENTS Page I Introduction 1 A Origin of Synpalpi 1 B Statement of Problem 1 II Description of Mutant Stock under Investigation 2 III Experimental Study 3 A Methods of Culture 3 B Stocks 5 G Grosses 7 1. Behaviour of mutant i n stock culture 7 2« Preliminary investigation of type of inheritance 8 (i) Crosses set up and results 6 ( i i ) Discussion of results 19 3* Temperature treatments 24 (i) Cold treatments 25 ( i i ) Heat treatments 45 ( i l l ) Discussion 57 4» Investigation of the lethality of the mutant 58 (i) Grosses of Zple and Xple synpalpi 58 ( i i ) C1B method 61 ( i l l ) Discussion 64 5» Determination of the linkage group 65 (i) Chromosome I 65 ( i i ) Chromosomes II and III 68 Page ( i i i ) Chromosome II 69 (iv) Chromosome III 70 (v) Summary 73 17 Discussion of Results 74 V Summary 78 71 Literature Cited 79 711 Plates 81 A B S T R A C T COLE, KATHLEEN MARGARET: THE STUDY OF A POSSIBLE NEW MUTATION, SYNPALPI, OCCURRING IN DROSOPHILA MELANOGASTER. The new mutation, synpalpi, which was found in the Xple Stock obtained from the Biological Supply House, Chicago, is so named because of the fusion of th£ antennae along the midline, a l l other Xple characteristics being normal except for a slight pro-trusion of the forehead. In stock cultures the mutant appears in greater numbers from normal Xple crosses than from synpalpi Xple crosses - with a predominance of female synpalpi. Experiments were carried out in order to determine the nature of the mutation, the type of inheritance involved, and the linkage group with which the mutation is associated. The mutant is not a gene mutation but a chromosomal aber-ration since Mendelian ratios were not observed to occur and since the mutation appeared irregularly but persistantly. A lethal factor is suggested by the small number of f l i e s resulting from synpalpi crosses; the absence or small number of synpalpi in the F of syn-palpi crosses but i t s reappearance in 2 - Qf> of the F progeny; and by the death of f l i e s in two different stages of development -early and late pupal stages. The association of the lethal factor with the "X" chromosome is assumed because few male synpalpil emerge. A deficiency of a part of the "X" chromosome would explain the death of the males and homozygous females. The possibility of a deficiency of a few bands at the tip of the T,Xn chromosome was sub-stantiated by a review of the literature on deficient "X" chromosomes. It is believed that synpalpi f l i e s are those deficient f l i e s which do manage to survive and that the fused antennae are caused by the move-- 2 -ments of the f l y and f o r c i n g of f l u i d to the head i n i t s e f f o r t s to e x t r i c a t e i t s e l f from the pupa case ( i t s tracheae being attached p o s t e r i o r l y to the c a s e ) . The Xple stock i s very s e n s i t i v e to v a r i a t i o n s from the o o © optimum temperature of 25 C. Temperatures of 35 C., 29 C , and 15 C. increased the l e t h a l e f f e c t of the d e f i c i e n c y - no synpalpi appearing i n the F or F of temperature-treated c u l t u r e s . be In the t e s t f o r l i n k a g e , the mutant appears to l-fce assoc-i a t e d with chromosomes, I and I I I - t h e r e f o r e the mutant i s not a simple d e f i c i e n c y of the "X" chromosome alone but appears to i n v o l v e the t h i r d chromosome as w e l l . The Study of a Possible New Mutation, Synpalpi, Occurring in Drosophila Melanogaster I Introduction A. Origin of Synpalpi On January 28, 1944, a culture of the multiple stock, Xple, was received, in a v i a l , by Mrs. Bath Fields Brink from the General Bio -logioal Supply House, Chicago. The v i a l contained several larvae which were placed in a culture bottle. Six f l i e s emerged showing Xple char-acters. These f l i e s were mated interse and on February 23 a mutation appeared in which the basal parts of the antennae were fused on the mid l i n e . This mutation s t i l l appears with great persistence i n the stock. oo 0 oo B. Statement of Problem The object of this study i s to determine the nature of the mutation, the type of inheritance involved, and the linkage group with whioh the mutation i s associated. oo 0 oo - 2 -II Description of Mutant Stock under Investigation The mutant stock under investigation i s in most respects similar to that of the normal Xple stock - the size, colour, and wings being identical. The distinguishing feature of the mutant i s the fused antennae , appropriately named "synpalpi", which are connected except ' at the immediate base and project out along the mid line of the head. The aristae, attaohed to the antennae, also protrude vertically as compared to the horizontal aristae of the normal Xple f l y . The head has a slight protrusion at the mid point, which i s probably caused by the pulling of the antennae when they fuse. Comparisons of normal and synpalpi f l i e s , dorsal and ventral views to illustrate these points, are shown in Plates I, I I , I I I , IV. An important observation i s the fact that the antennae are not fused imnediately at the base because i t has been noted that the antennae may separate and maintain the original protruding position. This mutation i s very easily recognized as seen in the plates named and in a lateral view the antennae may be seen to protrude, straight out from the head as compared to the normal antennae whioh curve downward ^  Plates WL • In some of the stocks the abdomen of the female mutants is abnormally pointed and appears squeezed. The significance of this w i l l be mentioned i n a later discussion. I l l Experimental Study A. Methods of Culture The chief culture requirements of Drosophila, oommonly called the fruit f l y , were supplied as follows$ adequate food conditions, livi n g yeast c e l l s whioh form the chief food of the larvae, an optimum temperature of about 25° C , and sufficient moisture. The adequate food conditions were met by a culture medium whioh provided a good physical substrate and a sufficient supply of sugars for good growth of yeast. The recipe for the medium i s given below. 1000 c c . water - bring to a b o i l 10 g. agar - add to above - b o i l 95 c.c. molasses) ) add to above - mix - b o i l 150 c.c. water ) 100 g. cornmeal - add to above - mix - b o i l several minutes. The medium was poured into culture bottles, previously corked with cotton and sterilised, u n t i l they were approximately one-quarter f u l l . Vfliile the medium was s t i l l hot, small pieces of paper towelling were placed in the bottles to provide a place for deposition of eggs and a resting place for larvae to pupate. If the cultures were to be kept for a time Moldex-A was added to eliminate mold. This was used in concentrates of 1 gram in 100 c.c. of 95 per cent alcohol. Five c.c.'s of this solution were added to the medium five minutes before fi n a l boiling was completed. Moldex -A was obtained from Glyco-Products Co., 148 Lafayette St., New York City. - 4 -The second condition, l i v i n g yeast c e l l s , was provided "by mix-ing a cake of Fleischmann* s Yeast in a dropping 'bottle of water and adding two drops of this suspension to the food surface in each culture bottle. The yeast was not sown u n t i l the bottles were to be used. Culture bottles were stored in the refrigerator therefore i t was neces-sary to allow them to reach room temperature before yeasting. The optimum temperature of about 25° C. was maintained by placing cultures in a thermostatically controlled incubator. The oultures were kept sufficiently moist by placing a pan of water in the incubator. Flies to be mated were added to the yeasted culture bottles. Since these f l i e s were under ether, in order to insure that they would not f a l l on the medium and so damage their wings or not be able to detaoh themselves, they were brushed into a paper scoop and care-fu l l y placed on the paper in the bottle. Etherization of the f l i e s was carried out by.shaking them from their culture bottle into a bottle, the a i r of whioh was saturated with ether. Such a bottle was called an etherizing bottle and was fitted with a cork which had a piece of cotton attached. To saturate the a i r of the bottle, ether was poured on the cotton and the cork f i t t e d tightly into the bottle. The f l i e s were removed as soon as a l l move-ments ceased - overetherization being avoided - especially i f the f l i e s were to be used for establishing cultures. Virgin females were secured from stock cultures by f i r s t shaking off a l l f l i e s . The females which emerged 12 r 15 hours later were virgin. B. Stocks The following descriptions of mutant stocks used were obtained from the hook by Bridges and Brehme (1944). The mutant stock under investigation, Synpalpi, and the stock from whioh i t arose are Zple. Xple - The characters soute (I - 0.0 4-), echinus (I - 5.5), cut (I - 20.0), vermilion (I - 33.0), garnet (I - 44.4), forked (I - 56.7) are present. Synpalpi also appears i n orange Xple stock. This stock i s similar to Xple - but the eye color, instead of being a complex of vermilion and garnet, i s orange. Orange = cd 2 - cardinal - formerly called orange. Appeared i n chromosome carrying non-lethal Ins. (2L 4-2R)P. Eye color yellowish vermilion - Ocelli white. BKjA. ' The stock used to test l e t h a l i t y on Chromosome I was C1B stock, which i s described more fu l l y later. The stock used to test for linkage group, Chromosome I, was yellow: y: yellow - Wallace, 1 - 0.0 Body color rich yellow - hairs and bristles brown with yellow t i p s . Wing hairs and veins yellow. Locus between IA5 and 1A8. Larval setae and mouth parts yellow to brown. Yellow occurs frequently by mutation. EK]_. - 6 -The stock used to test for linkage group, Chromosomes II and III, was Star dichaete: S/Cy D3/ln(3L)PMe. S: Star - Bridges, 2 - 1.3. Eye somewhat smaller and narrower than wild type and rough in texture from rounded, irregularly placed facets. Slight gleam from rumpled hairs on eye surface* Homozygote is lethal, dying in late embryonic stage* EK^  D3: Dichaete3 - Plunkett, 3* Inseparable from In(3L)P. Wings extended and lifted at 45 degrees from the 6 body axis and elevated 30 degrees above* Alula gone* Viability of D3/+ is better than that of T)/4-. D3/D and D3/p3 are lethal* EEgA. Cy: Curly - Ward, 2, Associated with In(2L)0y. Wings curled up strongly, rarely overlapping wild type at 25°C. Locus within inversion. Homozygote is lethal. BKgA. In(3L)P: Inversion (3L) Payne - Payne. In third ohromosome - lethal in homozygotes. > \ Little C.O. with Me. Me: Moire - Muller, 1929. 3 - 20 ± Eye has watered-silk shimmering iridescent pattern, due to ring of 6 flecks around normal fleck. Eye color transluoent and brownish with little color in outerlayers and interior. Homozygous lethal. RK,A. The Btook used to test for linkage group Chromosome II, was Lobe curly: L: Lobe-Bridges, 2 - 27,0. Heterozygous L eye reduced in size with nick in anterior edge and lower half of eye reduced more than upper* Homozygous L eye much smaller and less viable. Best used as recessive, RK^ . Cy: Curly - described previously. The stock used to test for linkage group, Chromosome III, was Dichaete glued: 3 3 ' D : Diohaete -described previously. Gl: Glued - Ives, 3 - 41,4. Eyes smaller and oblong with facets rounded, surface smooth and shiny. Homozygous lethal. RK^ . The wild stock used was Urbana-S. Urbana-S (Urbana-Special): A stock selected by Bridges from one oolleoted at Urbana, Illinois. The body color is somewhat lighter than the standard wild type. The salivary chromosomes were found by Bridges to be normal. (Description received from Schultz). oo 0 oo C. CROSSES 1, Behaviour of mutant in stock culture. The mutant, Synpalpi, appears persistently in the Xple and Xple orange stock cultures. In the Xple stock it has been noted that there is a greater predominance of female synpalpi flies - very few male synpalpi appearing. In the Xple orange stock more male synpalpi ap-pear hut they are st i l l in the minority. Fewer synpalpi appear in stocks whioh are set up using only synpalpi males and females than in those set up using normal Xple flies. Many flies appear in these stocks whioh have their antennae protruding straight out but not connect ed -as described on Page 2. Some stocks in the larval stage whioh suf-fered a slight decrease in temperature for two days, due to unusually cold weather* contained no synpalpi flies for six days after emergence of flies began. 2. Preliminary investigation of type of Inheritance. The first, step in this investigation was to try to obtain a homozygous mutant stock and a picture of the inheritance pattern. Various crosses were set up in order to achieve these aims. (i) Crosses set up and results. Cross #1 Bottle #2: Cross of 2 Xple orange synpalpi5°x 3 Xple orange synpalpi Cr^o^. P 1 results are listed in Table I. Table! B #2 Non sal ? ? Synpalp 1 ?? Nov. 4 5 6 7 Total 15 12 2 1 22 12 4 1 (died) 1 30 38 1 1 The original parents from Bottle #2 were transferred to Bottle #20 -E 1 count consisted of 5 normal errand 5 normal?.?. The total F 1 count was therefore: 35 normal Xple orange cTo-? 42 normal Xple orange ? ?, 1 synpalpi Xple orange C? , and 1 synpalpi Xple orange ? . F 1 interses were set up - Bottles #21, 22, and 201. In Bottle #23 a cross was set up of the F 1 synpalpi ? z F* normal o \ F 2 results are listed in Table II. The abbreviation B for Bottle will be used hereafter. Table II Norn sal ? ? Sym >alpi ?9 Non nal 99 Synp alpi Norn a V ial B #21 Nov. 15 20 22 Total 2 3 4 6 4 8 2 1 B #22 Nov. 15 18 20 22 23 Total 1 5 4 1 2 7 3 3 2 1 2 B #23 Nov.- 18 20 22 Total 3 2 1 3 4 2 9 16 0 3 6 9 10 16 0 5 B #20 Nov. 22 25 26 27 Total 12 5 14 18 13 10 10 19 2 4 1 Totals of a l l Fl interses 68 86 6 9 49 52 • 6 1 Since length of life histories varied an accurate account was kept of the time each stage was reached. This is shown in Table III -giving times for appearance of F 1 and F 2. TABLE III BOttle NO. Set Up Eggs Small Larvae Pupae First F 1 or F 2 Total Length in Days 2 Oot. 24 Oot. 26. Oot. 27 Oot. 30 Nov. 2 9 20 Oot, 30 NOV. 1 Nov. 2 Nov. 5 Nov. 8 9 21 Nov. 4 Nov. 5 Nov. 6 Nov. 10 Nov. 13 9 22 Nov. 5 Nov. 6 Nov. 8 Nov. 12 Nov. 15 10 20l Nov. 9 Nov. 10 Nov. 12 Nov. 16 Nov. 18 9 23 Nov. 5 Nov. 6 Nov. 9 Nov. 13 Nov. 17 12 -10-In a cross of synpalpi o 3 x synpalpi ? one would expect to get a high percentage of synpalpi F 2 - but the results showed only approximately 9% synpalpi in F 2. It is noticeable in the results in Table II that in both normal and synpalpi F 2 the males were not so plentiful as females. Both of these latter statements may be ac-counted for by a lethality of this synpalpi mutant - since the male has only one X chromosome. The F 2 results from the cross of P 1 synpalpi $ x normal also seemi to indioate a lethality character of the mutant -? since there was a very low count - and no synpalpi appeared. There was also a prolonged life history of 12 days - as compared to the 9 and 10 day duration of the others, Table III. Gross #2 B #3: Cross of 2 Xple orange synpalpi ?? x 3 Xple orange oV. F 1 results are listed in Table 17 - along with F 1 results of B #30 -parents from B #3. TABLE IV B #3 Nor CrV ooal ?? Synj >alpi ?¥ B #30 Non o V ooal ?? Synj o V >alpi ?* Nov. 4 5 6 7 Total 29 9 1 3 23 7 4 1 1 Nov. 9 12 13 15 Total 3 10 6 4 4 7 1 4 2 42 35 0 1 23 16 0 2 Total F 1 Count 65 51 0 3 F* interses were set up - B #31, 32, 301, 303. F 1 synpalpi crosses* B #33 9 1 P 1 synpalpi $ x 2 F^ c?if B #302» 2 F 1 synpalpi 5 £ c / V - 11 -F 2 results are li s t e d in Table 7. Table 7 Hon nal ?? Synp alpi ?? Nor coal 2? Synp a l p i ¥9 Non cry sal 99 Synpi i l p i B #31 HOT.13 15 18 20 22 Total 1 3 3 4 2 1 3 1 4 3 1 B #32 Nov*18 20 22 25 Total 12 6 6 1 11 7 2 3 B #302 Nov*25 27 29 Total 4 8 5 5 17 5 17 27 0 0 B #33 NOV*18 20 22 25 Total 2 14 10 3 4 11 13 5 1 2 13 12 1 0 25 23 0 0 B fSO 1 Nov*18 20 22 25 27 Total 6 5 1 30 21 3 2, 10. 18 16 3 1 1 2 1 1 B #303 Nov.25 27 29 Seo. 2 Total 3 4 3 2 2 5 2 1 1 1 29 33 3 0 Total P l Interses 111 95 7 6 63 49 4 5 10 11 2 1 Length of l i f e histories are given in Table 71 Table 71 Bottle No* Set Up Eggs Small Larvae Pupae Fir s t _ F 1 or" F 8 Total length In Days* 3 Oot.22 Oot.24 Oct.25 Oot.28 NOV* 1 9 30 Oot. 28 Oot.29 Nov. 2 Nov. 6 Nov. 8 11 31 Nov. .4 Nov. 5 Nov. 6 Nov. 9 Nov.13 9 32, Nov* 5 Nov. 6 Nov. 8 Nov. 12 NOV.13 8 30 1 Nov* 9 Nov.10 Nov. 12 Nov.16 Nov*18 9 30 3 Nov.15 Nov. 16 Nov.18 Nov.22 Nov.25 10 33 Nov. 5 Nov. 6 Nov. 8 Nov.15 Nov.18 13 30 2 Nov.12 Nov.13 Nov.15 NOV.20 Nov*25 13 In this oross of Synpalpi? x normalcfithere was a more balanced number of males and females* Synpalpi appearing as a result of F 1 i n -terse constituted only 6% of the total F 2. The cross of F 1 synpalpi $ x.P 1 normal o^yielded only 3$ synpalpi - a l l males. There was also a very poor ratio of males and females* A comparison of Tables III and 71 shows that the life history of crosses of F 1 synpalpi? x normal a71 increased in duration - being 12 to 13 days from date set up to date of appearance of first F 2 flies. Also apparent in Crosses #1 and #2 was the fact that in general most synpalpi flies appeared on the third or fourth day after appearanoe of first F 2 flies. Cross #3 - Reciprocal of #2. B #5: Cross of 5 Xple orange $?x 2 Xple orange synpalpicr^ F 1 results are listed in Table, 711 • Table 711 B #5 Normal Synpalpi o V ?? Nov. 8 1 4 9 3 3 12 1 3 13 1 1 Total 5 11 1 0 An •F^  interse was set up in B #51. F**" synpalpi cross: B #52 - F 1 synpalpio^x F1. normal? • F 2 results are listed in Table T i l l . Table T i l l B #51 Nor o^o^l mal ? $ Synp* ilpi '?* B #52 Norr cry tttl 1* Synpi alpl 9-9 NOV. 20 22 25 27 Total 30 5 11 27 12 8 1 1 1 1 N0V.27 29 Deo. 2 Total 11 2 6 3 1 46 47 0 4 13 8 1 0 Length of life histories are given in Table IX. Table IX Bottle No. Set Up Eggs Small Larvae Pupae First F 1 or F 2 Total Length In Days 5 Oot.30 Oct.31 -Nov. 2 Nov. 5 Novi 8 9 51 Nov. 8 Nov. 9 Nov.13' Nov.18 Nov.20 12 52 Nov.13 Nov.14 Nov.17 Nov.21 Nov.27 14 The F 1 count (Table VII) was very low considering that five females were used. Males constitute only one-third of the total -normally should be one-half. Both these facts suggest again the lethal-i t y of the mutant synpalpi. The F 2 count (Table VIII) - result of F 1 interse - showed only 4$ synpalpi - a l l female. Cross #2 - reciprocal of this - gave 6% synpalpi - but both male and female appeared. As in previous crosses the F 2 oount - result of F 1 synpalpi x F*" normal - was very low and the l i f e history, very long - (14 days). Cross #4 B #4 - Cross of 2 wild$?x 2 Xple orange synpalpi Oo^. F 1 results are l i s t e d i n Table X. A l l F 1 f l i e s were wild type. Table X B #4 Non d V nal ? ? Synpa o V l p i ? ? Nov. 9 12 13 15 Total 5 13 4 3 9 5 1 2 25 17 0 0 An F 1 interse was set up - B #41 and #42. F 2 results are l i s t e d in Table XI. - 14 -Table XI WiM Xple Orange Nor mal ?? Synpj oV"> i l p i $? Non o V aal ?<? Synp o V alpi ¥$ B #41 Nov. 22 25 27 29 Total 1 13 1 1 1 10 1 4 4 1 16 19 0 0 1 0 0 0 B #42 Nov. 25 27 29 Total 18 10 8 19 13 12 1 2 2 36 44 0 0 5 0 0 0 Total of F^- interse 52 63 0 0 6 0 0 0 Length of l i f e histories are given i n Table XII. Table XII Bottle Set Up Eggs Small Pupae First F 1 Total Length NO. Larvae or F 2 In Days 4 Oot.30 Nov. 1 Nov. 3 Nov. 6 Nov. 8 9 41 Nov. 9 Nov.10 Nov.13 Nov.16 Nov.21 12 42 Nov.12 Nov.13 Nov.15 Nov.20 * Nov.23 11 No synpalpi appeared either in F* or F 2. Cross #5 -Reciprocal of #4 B #6 - Cross of 1 Xple orange' synpalpi + x 4 wild^ty 7. F 1 results are li s t e d in Table XIII. A l l F 1 f l i e s were wild type. Table XIII B #6 Non ^ 9 i Ipi Nov.15 18 Total 6 4 6 6 10 0 0 - 15 -P 1 interses were set up - B #61 and #62. P 2 results are l i s t e d in Table XIV. Table XIV Wild Xt>le Oranse Non nal ? ? Synp alpi ?* Nor orVl mal Synp alpi <?$ B #61 Nov.29 Dec. 2 4 Total . 4 4 3 3 4 1 11 8 0 0 0 0 0 0 B #62 Nov.27 Dec. 2 4 Total 8 5 1 9 6 2. 1 14 17 0 0 1 0 0 0 Total of F! interse 25 25 0 0 1 0 0 0 Length of l i f e histories are given in Table XV Bottle No. Set Up Eggs Small Larvae Papa© First F 1 or F 2 Total Length In Days 6 Nov. 5 Nov. 6 Nov. 8 Nov.11 Nov.14 9 61 Nov. 15 Nov.16 Nov.18 Nov.25 Nov.29 14 62 Nov.15 Hov.16 Nov.19 Nov.24 Nov.27 12 Cross #4 and i t s reciprocal, #5, had extremely low F* counts -as seen in Tables X and XIII - no synpalpi appearing. Cross #4 P* fe-males were i n the minority as compared to Cross #5 where the males were in the minority. The F 2 counts of both crosses - Tables XI and XIV -were also very low, Cross #5 having the lowest. No synpalpi appeared in - 16 -P 2 progeny in either G r o s s - this may he explained by low counts and lethality of mutant. Cross #5 which had the lower F 2 count also had longer l i f e histories of F 1 interse crosses - as seen in Tables XII and XV. Cross #6 B #300 - Cross of 2 Xple synpalpi ??x 4 Xple synpalpicr^ 1 F results are l i s t e d in Table XVI. Table XVI B #300 Nor or>cr coal 99 Synps il p i 99 Nov. 9 12 13 Total 3 10 8 1 13 6 21 20 0 0 F interses were set up - B #301 and #302. F 2 results are li s t e d in Table XVII. Table XVII B #301 Hor • cr?cr] coal 99 Synp alpi 92 B #302 Hon nal 9? Synp a l p i Nov.22 25 27 29 Total 1 15 3 6 4 8 5 3 1 1 1 1 Kov.25 27 ' 29 Dec. 2 Total 6 6 2 4 6 9 1 7 2 1 1 25 20 1 3 18 23 3 1 Total of pl interse 43 43 4 4 Length of l i f e histories are given in Table XVIII. - 17 -Table XVIII Bottle Ho. Set Up Sggs Small Larvae Pupae First gF 1 or W Total Length In Days 300 Oot.30 Nov. 1 Nov. 3 Nov. 6 Nov. 8 9 301 Nov. 9 Nov.10 Nov.12 Nov.18 Nov.21 12 302 Nov.13 Nov. 14 Nov.16 Nov.20 Nov.23 10 Comparing FA results of Cross #1 (Table I) Xple orange - the F 1 count in this cross (Table 2SVT) was lower and had no synpalpi as Cross #1 had. The F 2 count showed 9fa synpalpi - the same as that of Cross #1. Length of l i f e histories of Cross #1 (Table III) and Cross #6 (Table XVIII) were also similar. Cross #7 , B #100 - Cross of 2 Xple synpalpi 5 Xple o ^ , F 1 results are l i s t e d in Table XIX. Table XIX B #100 Non tnal 9? Synj o-"Vr> >alpi ! 9? Oct. 25 28 Total 17 13 17 13 30 30 0 0 F 1 interses were set up - B #101 and #102. F 2 results are l i s t e d in Table XX. Table XX o B #101 Norn a l 9 2 5 $ P " B #102 Nov. 4 5 6 7 9 12 13 Total 2 6 1 2 2 1 1 2 1 2 1 1 Nov. 6 7 8 12 13 Total 3 8 14 4 1 2 10 18 6 l i 2 1 29 37 0 5 Total of F l interse 40 46 1 6 11 9 1 1 Length of life histories are given in Tattle XXI. Table XXI Bottle HO.. Set Up Eggs Small Larvae Pupae First F 1 or F 2 Total Length In Days 100 Oct.11 Oct.13 Oct.16 Oct.19 Oct.22 11 101 Oct.25 Oct.26 0ct.28 HOV. 1 Nov. 4 10 102 Oct.28 Oct.29 Oct.31 Nov. 3 Nov. 6 9 Synpalpi occurring in F 2 constituted 8% of total - higher than that of the similar cross of Xple orange (Cross #2). In Cross #2 synpalpi were males - as contrasted to this cross, where synpalpi were mostly females. Cross #8 - Reciprocal of #7* B #200 - Cro ss of 1 Xple ? x 1 Xple synpalpi cr57. F 1 results are listed in Table XXII. Table XXII B #200 Normal Synpalpi ?9 cr>cr> 99 Nov. 8 2 3 9 6 5 12 6 5 1 13 6 7 1 15 5 6 1 ' 1 Total 25 31 3 1 F 1 interses were set up - B #201, #202. Crosses were also set up of F 1 synpalpi'. B #203 = 2F1 synpalpi cf^k P* normal ? . B #204 s F 1 synpalpi o/* x F 1 synpalpi ? - died. F 2 results are listed in Table XXIII. - 19 -Table XXIII B #201 Hon c/M nal S? Synp oVj alpi ?? B #202 Nor oVI ml %? Synp alpi 99 B #203 Nor o V | mal Syn] jalpi f ?? HOT,22 25 27 Total 21 13 2 18 9 2 4 Nov,22 25 27 29 Total 2 11 30 12 6 7 31 9 1 3 1 Nov.25 27 29 Deo* 2 Total 4 11 9 7 9 3 15 4 1 36 29 4 55 53 1 4 31 31 1 Total of a l l F 1 interses 91 82 1 8 Length of l i f e histories are given i n Table XXIV* Table XXIV Bottle NO. Set Up Eggs Small Larvae Pupae F i r s t F 1 Of F 2 Total Length In Days 200 Oat.28 Oct .30 Nov. 1 Nov. 4 Nov. 8 11 201 NOV. 8 Nov. 9 Nov.13 Hov.18 Nov.21 13 202 Nov.13 Nov.14 Nov.16 Nov.18 NOV.21 8 203 Nov.13 • Nov .14 Nov.16 Nov.19 Nov.22 9 In the F 2 oount, synpalpi constituted 5$ of the t o t a l . The Bimilar cross of Xple orange (Gross #3) gave 4% synpalpi - which i s very near that of this cross (#8). Another similarity of these two crosses i s the predominance of female synpalpi. N.B. In a l l tables normal refers to normal antennae as opposed to fused antennae (synpalpi)* ( i i ) Discussion of results. The results of the preceding crosses have been summarized and tabulated in Table XXV. The time factor for the emergence of F 1 f l i e s and F 1 counts cannot be considered significant since the crosses were made at d i f -ferent times, many of the developing f l i e s being at a c r i t i c a l stage - 20 -over the weekend when the temperature dropped in the incubators, and since different numbers of parents were used in the various crosses* In general, though, the low iP" and F 2 counts suggest a low fert i l i ty of the mutant synpalpi. Crosses #1 and #6, whioh were crosses of synpalpi 9 x synpalpi a produced 9* synpalpi in the F 2 - synpalpi appearing in males and females in equal numbers. Few or no synpalpi were produced in the F* - a fact whioh suggests that synpalpi is not a simple dominant* Another i n -dication that the mutant is not a dominant is seen in the crosses of wild x Xple orange synpalpi (Grosses #4 and #5), where no synpalpi a p p e n d in the P 1 „ the In Crosses #2 and #7 there were slightly fewer synpalpi in the F 2 than in crosses #1 and #6, In the case of Cross #2* Xple orange synpalpi ? x Xple orange ° * * where 2,5$ synpalpi were produced in the F** there was a fairly equal number of synpalpi F 2 males and females* In the case of Cross #7, Xple synpalpi? x X p l e O ' 7 , where no synpalpi were produced in the F 1 , there was a predominance of female synpalpi in the F 2. In crosses #3 and #8, where the mutation was introduced by the male, there were fewer synpalpi in the F 2 than there were in Crosses #2* #7, #1 or #6* - the female synpalpi flies being in greatest numbers* These crosses (3 and 8) also produced 6% synpalpi in the F 1 with a greater predominance of male synpalpi* The fact that fewer synpalpi appeared in the T1 than in the F 2 may indicate that most homozygotes die in the F 1 and the heterozygotes, by segregation and recombination, produce more synpalpi in the F 2 . In o - 21 -general there is no Mendelian segregation, but there does appear to be a hereditary basis to the mutation sinoe chance could not account for al l the similarities in ratios obtained* Summary of the Preliminary Investigation* dross No, Constitution No, of pJ- Syn-palpi F 1 NOo of pl..Sjn-palpi 9? No. of P xSyn-palpi Average Length of l i f e history of F 1 interse No. of P 2 % Syn-No. of P l Syn-palpi 99 No. of p l Syn-palpi 1 2 Xple orange synpalpi' x 3 Xple orange synpalpic - 79 2.5 1 1 10 days Interse: I69 74 95 9 p l Syn<? ) l t r x: P l <r> ) 0 6 cf> s 9 9 0 9 0 6 0 2 2 Xple orange synpalpi^ x 3 Xple orange cr> 119 65o">54? 2.5 3: 0 9 " Interse: 219 118cr>l01? pl Syn? ) 1 Q q 49 cr1 60$ 6 ,-• 3 6 0 7. 3 35 5 Xple orange ? x 2 Xple orange synpalpi* 17 ^6o-7ll9 6.0 0 1 12 '» Interse: 97 46o-->51? F 1 ? * ) 2 2 P 1 Syn a*) 14. <r> 8 9 4 5 4 0 0 1 4 .2 wild 9 . x 2 Xple orange synpalpi 42 ^(wild) 25o^.7? 0 0 0 11.5 11 Interse: 121 (Wild: 520*63?) (Xple orange 6oj 0 0 0 5 1 Xple orange synpalpi x. 4 wilder* R 16 (wild) 6cr>10<? 0 0 0 13 " Interse: 51" (nmd:25<^259; (Xple orange lcr? 0 0 0 TABLE XXV (Continued) Cross No. Constitution No. of pi -i Syn-palpi p i No. of F x Syn-palpi 9? NO e Of P 1 Syn-• palpi cr>cf> Average Length of l i f e history of F l Interse No. of P 2 Syn-No. of p i Syn-palpi 99 No. of pi Syn-paTji o-ly 6 2 Xple synpalpi^ x 4 Xple synpalpio 41 => 210^20$ 0 , 0 0 11 days Interse: 94 47 ori 47 9 9 4 4 7 2 Xple synpalpi? x 5 Xple synpalpio 60 , 0 0 0 9 .5 "* Interse: 93 41 o--7 52 ? 8 6 1 8 1 Xple9 x 1 Xple synpalpio 60 ^ 280^32? 6o0 1 35 11 "L Interse: 182 92 cr> 90 9 pi 9 ) x P 1 cr) ) 63 Syn ) 31 cf1 32 9 5 2 8 1 1 0 o 3, Temperature treatments* Following the preliminary investigation of the type of in-heritance an attempt was made to increase the penetrance of the mutation by prolonged cold and heat treatments* Villee (1942 and 1944) des-cribed the prooess by which the mutant arlstapedia, in whioh the arista is a tarsus with bristles and claws, was affooted by temperature* Since this mutant arlstapedia also affects the antennae, it was thought that the experimental methods used by Villee oould be applied to the mutation synpalpi* In Villee*s cold treatments a temperature of 14*4° 0. and an age of four days of development (from the egg) proved most sensitive, as the cold decreases the velocity of development and enables the mutant gene to work at a lower rate for a longer time, thus producing an effect. Heat treatments of 29 - 35° C* temperature were used whioh decreased the expression of the mutant arlstapedia. Balkashira (Villee 1944) reported that in arlstapedia seg-mentation of the antennae disk is begun in the two day larvae along with the leg diso* In normal flies it occurs in four to four and one?-half day larvae. Braun (1940) also reported this and remarked that the mutant arlstapedia acts by shifting the initiation of the embryologioal prooess to an earlier point in development* Robertson (193?) stated that the head is formed at the end of the prepupal period and the antennae develop immediately following for 66 hours* Considering al l this it was decided to subject larvae of ages 3 - 4, 4 - 5, and 5-6 days of development to heat and cold treatments, as these ages should include the sensitive period of development of the antennae. - 25 m (i) Cold treatments. A temperature of 15° C. was used in all cold treatments. On December 18 at 1:15 p.m. the following orosses were set up ^* al l females used were virgins; #1 - 13 Xple? x 15 Xplectf) obtained from Xple } synpalpi stock bottles #2 - 14 Xple? x 15 Xple cf) #3 - 1 Xple synpalpi $ x 3 Xple snypalpi cr1 #5 - 5 Xple orange ? x 10 Xple orangecf) obtained from Xpls ) orange synpalpi #6-5 Xple orange ° x 11 Xple orange cr) stock bottles #7 - 2 Xple orange synpalpi? x 4 Xple orange synpalpi cr' On December 19 at 1:15 p*m. the parents were transferred to new .bottles: #1 to #10, #2 to #20, #3 to #30, #5 to #50, #6 to #60, #7 to #70. On December 20 at 1:15 p.m. the parents were transferred again-to new bottles: #10 to #100, #20 to #200, #30 to #300, #50 to #500, #60 to #600, #70-to #700. On December 23 at 1:15 p.m. the parents were removed from bottles #100, 200, 300, 500, 600 and 700. Bottles #1, 2, 3, 5, 6, 7 -age 4-5 days and bottles #10, 20, 30, 50, 60, 70 - age 3-4 days <* were placed in the 15?C. chamber. Bottles #100, 200, 300, 500, 600, 700 were used as controls and were kept at the normal temperature of 25° C. Bottles were removed from the cold on January 13. First F- flies of controls emerged on January 2. - 26 -Of the bottles placed In the cold #3, #30, and #5 produced no larvae and #1, #10 and #20 were the only ones to produce living F 1 flies. F* counts of bottles placed in cold are listed in Table XXVT. Table XX7I - : I .  #1 Non mal ? $ Synpe cry ilpi #10 Nor cry raal 99 Synpa cry Ipi 9? #20 Nor °V| mal ?? Synp! alpi 99 Jan.20 27 Total 1 1? "Jan. 27 Feb. 3 35 1 32 2 Jan.27 29 Feb. 3 32 8 5 26 7 12. 1 1? 0 0 Total 36 34 0 0 Total 45 45 0 0 F 1 interses were set up as follows: IF 1? ? x IF 1 - Jan. 27 - died SF^?? x 3F1 Jan. 27 3F1 9 9 x 3F1 o-V7- Jan. 27 3F1 9? x 3F1 Jan. 29 F 2 oounts of the above F 1 interses are listed in Table XXVII. Table XXVII f i o 1 Norms i l 9? Synps o-V ilpi <?? #201 Non cry nal 9? Synps ilpi 99 #202 Nor mal ! o_? Synp talpi 99 Feb. 10 11 12 Total 33 22 11 39 29 11 Feb. 10 11 12 Total 9 20 19 15 31 12 l Feb. 10 11 12 4 6 11 9 15 10 66 79 0 0 48 58 0 l Total 21 34 0 0 Bottles #2, #50, #6, #60. #7, #70 produced no living adult flies - a l l died in the pupal stage. In order to determine the state at whioh death occurred and the sex of the flies, the pupa cases were opened and examined under the binocular. A petrl dish filled with wax and two fine needles proved most #1 : B #1 : #10t B #101 : #20: B #20X : B #202 : - 2 7 -satis factory for dissection. The terms "larval" state and "adult or emergent" stage are used in describing the flies in the pupa oases. "Larval" stage refers to the stage at whioh the body segmentation occurs. The body which is yellow-shite, is very soft and filled with a fluid. The divisions of head, thorax and abdomen are evident - the wings show as projecting, narrow balloon-like flaps from the thorax and the eye regions are slight bumps on the head*, Fig* 1. Experimentation and examination in-dicated this as the 6 day old period in development* "Adult" or "Emergent" stage refers to the period just before emergence* The wings, body, legs and eyes are well developed. The antennae appear as dark protrusions from the head or are well developed with bristles. - 2 8 -Dissection of papa oases: B #1 No larvae v Sampling of 22 pupa oases : Emergent: 18 cryp3 synpalpi) 3 9£ Larval : 1 B#2 No larvae 44 pupae : Emergent: 34 oV'Z 99 Larval : 7 B #6 No larvae 30 pupae : Emergent: 21o&(i synpalpi) Larval : 9 B #7 No larvae 34 pupae : Emergent: 110^0^(1 synpalpi) 4?? Larval : 19 B #10 No larvae Sampling of 19 pupa oases : Emergent: 13 0 -^3 synpalpi) 5 99 Larval : 1 B #20 No larvae 28 pupae ; Emergent: 19 synpalpi) 2 ? 9 Larval : 7 B #50 No larvae 24 pupae : Emergent: 11 cr^S 94? Larval : 10 B #60 No larvae 31 pupae : Emergent: 11 c^Vl ? Larval ; 19 B #70 No larvae 41 pupae : Emergent: 12 cr^ 7 synpalpi) 3 99 Larval : 26 F counts of controls are listed in Table XXVIII. Table XXVIII #100 Normal Synpalpi #500 Normal Synpalpi #700 Non oal Synpalpi Op 9? crV 99 99 OCT | 99 99 9? Jan* *6 38 37 4 4 Jan. 6 6 3 3 3 Jan. 6 11 18 5 12 7 7 12 0 0 7 1 0 1 2 7 3 1 2 5 8 7 4 0 0 8 0 2 0 0 8 0 0 0 1 10 9 10 0 2 . 10 14 16 0 1 Total 61 63 4 6 Total 21 21 4 6 Total 14 19 7 18 #200 #600 Jan. 6 42 40 0 1 Jan. 6 20 31 2 7 7 11 15 0 0 7 4 4 0 2 8 7 7 2 0 8 1 0 0 1 10 3 4 0 0 10 2 0 0 0 Total 63 66 2 1 Total 27 35 2 10 B #300 - no eggs or larvae* orosses of controls were set up as follows: #100 : B #101 : F 1 interse of normal-39 x S d - January 6, B #102 : F 1 interse of synpalpi-2? x 2o->- January 8. #200 : B #201 : F * interse of normal-3? x 3 o-~> - January 6, B #202 : FT" interse of synpalpi-1? x 2cr>- January 8, #500 : B #501 : F 1 interse of normal-3? x 3 o"7 - January 6, B #502 : F 1 interse of synpalpi-29 x 2o*- January 7, #600 : B #601 : F 1 interse of normal-3?x 3 - January 6 B #602 : F 1 interse of synpalpi-3 9 x 2er>- January 7 #700 : B #701 : F 1 interse of normal-39* 3 or 1 - January 6 B #702 : F 1 interse of synpalpi- 3? x 3o-7- January 8 B #703 : F 1 synpalpi x F 1 normal- 39 x 3o-7 January 8 F 2 results of controls are listed in Table XXIX. - 30 -Table XXIX For nal Synp. oV alp: %9 Non o-V> nal 9? Synp o-Vi a l p i f¥ Nor o-7f mal Synp orV a l p i 99 #101 Jan.l? 20 21 Total 14 23 17 16 21 19 0 0 0 1 2 1 #501 Jan. 17 2C 21 19 23 17 23 21 15 0 1 0 3 1 7 #701 Jan.17 20 21 21 17 11 25 16 15 3 1 3 1 3 4 54 56 0 4 Total 59 59 1 11 Total 49 56 7 8 #102 Jan*20 21 24 Total 13 14 26 17 14 28 2 0 3 2 1 2 #502 Jan .20 21 24 12 17 21. 15 18 23 0 2 1 2 1 2 #702 Jan.20 21 24 22 10 21 27 11 17 2 0 4 3 1 5 53 59 5 5 Total 50 56 3 5 Total 53 55 6 9 #201 Jan.17 20 21 Total 23 37 9 27 35 10 0 0 0 3 2 1 #601 Jam. 17 20 21 21 13 21 23 14 20 1 0 0 4 2 2 #703 Jan.20 21 24 Total 13 17 21 16 16 20 1 0 0 0 0 0 69 72 0 6 Total 55 57 1 8 51 52 1 0 #202 Jan,20 21 24 Total 9 19 29 13 20 27 2 0 1 3 1 0 #602 Jan*20 21 24 18 16 21 17 19 17 1 2 1 0 3 3 57 60 3 4 Total 55 53 4 6 Additional oold treatments we're carried out in order to include the age group 5 - 6 days in the treatment. On January 24 at 12 a.m. the following crosses were set up: #1000 3 Xple synpalpi9?x 1 Xple synpalpi cr 7 #2000 12 Xple x 28 Xple o-V>(from Xple .synpalpi stock) . #3000 2 Xple orange synpalpi?& 3 Xple orange synpalpi o-V #4000 8 Xple orange ?£c 10 Xple orange o^from Xple orange synpalpi stock). The parents were removed January 25 at 12 a.m.. On January 30 at 12 |.m. the bottles were placed i n the cold -cultures at an age of 5 - 6 days. - 3 1 -The hot ties were removed from the cold on February 12, 1 No F f l i e s emerged, therefore, the pupa cases were opened and examined* B #1000 No larvae Sampling of 15 pupae : Emergent: . 9 6 9 Larval : 0 B #2000 No larvae Sampling of 15 pupae : Emergent: 5 7 $ Larval : 3 B #3000 No larvae Sampling of 15 pupae : Emergent: 4cr 78? Larval : 3 B #4000 No larvae Sampling of 15 pupae : Emergent: 10 0^3$ Larval : 2 The parents removed from bottles #1000, #2000, #3000, #4000 were placed in bottles #11, 2 1, 3 1, 4 1, respectively on January 27 at 5 p*m* Another transfer was made on January 28 at 5 p.m. to bottles used in heat treatment described later. The bottles were placed in the cold on January 31 at 5 p.m., when cultures were at the age of 3 - 4 days, and removed on February 14 at 5 p.m. F^ began to emerge on February 17. F 1 results are li s t e d i n Table XXX. - 3 2 -•Table XXX n 1 Normal Synpalpi &1 Normal Synpalpi °v ?? ? ? ? ? oV ?? Feb.17 3 9 Feb. 17 2 1) to 18 9 4 18 17 4 7 1) normal 19 1 19 53 27 20 0 0 20 1 5 1 5 Total 1 3 1 3 0 0 Total 8 5 9 1 0 2 normal IP 1 # 4 1 Feb.17 1 Feb. 17 1 l(cued) 18 1 18 8 19 19 4 20 2 0 Total 2 0 0 0 Total 1 3 (U 0 0 F 1 interses were set up: .#lj : 3 F1?? x 3 F W February 17 #2* : 3 F1?? x 3 F^Februsry. 1 ? #2g : 2 F 1 (synpalpi-* normal)?? x 3 FV C A February 18 P 2 results are tabulated in Table XXXI Table XXXI IT 1 ! Normal Synpalpi #2i Normal Synpalpi #2^ Normal Synpalpi J. o V ?? 9 ? •ia ?? 0 * 0 * & crV ?v Mar. 4 5 6 Mar.3 10 1 6 Mar.3 2 4 16 5 4 3 4 9 3 4 8 3 7 16 19 5 1 1 10 5 3 4 7 8 6 7 1 0 Total 2 5 28 0 0 Total 38 3 5 0 0 Total 36 2 3 0 0 The pupa cases containing dead flies were opened and examined. Observations. B $0- : No larvae 1 5 pupae : Emergent:' S c r V U - part way out of oase) ( 4 -. palps not developed) 1 0 ? ? ( 7 - young-palps not devel-oped. ( 3 - part way out of case) - 33 -In these f l i e s i t was noted that the front of the head protruded as in Fig. 2 - ventral view. Pig. 2 - Ventral view of head of f l y i n pupa oase, showing protrusion of head. B #21 No larvae A l l f l i e s emerged B l ^ 1 No larvae 2 pupae : Emergent: 2?? palps folded down i(•palps* refers to antennae) B #41 No larvae 94 pupae : Emergent: 61otb^ 21 - forehead bulged -palps protruding and well developed. 35 - palps folded -forehead not bulged - 3 of these show traoheae attached to posterior of pupa case. 1 - very young - head poorly formed - larval jaws attached. 2 - young - traoheae attached. 2 - emerged - dragged case. - 34 -30?? : 21 palps folded 6 palps protruding and forehead bulged, 1 - head not well developed - no pigment in eyes. 1 - no head 1 emerged - wings deformed. Larval : 3. Interesting notes on the dissection of pupa cases: 1* Plies whioh died when half emerged from the pupa oases had a silvery covering over their bodies which stuck the limbs and wings to-gether. There was a milky secretion from the posterior part of the body which was very sticky. The secretion is stated by Kaliss (1939) to be a substance containing calcium carbonate. 2. In some cases the traoheae of the larva were attached' to the pupa case making it impossible for the fly to detach itself and so emerge. 3. On applying pressure to the head the antennae of the fly, with folded antennae, protruded and tended to approach eaoh other and fuse. It would seem, then, that pressure on the head on emerging may cause synpalpi. The pressure may be caused by a deformed pupa oase or by contractions of the body of the fly in its efforts to detach the tracheae from the pupa case at the posterior. - 35 -Controls for bottles #1000, 2000, 3000, 4000 and #11, 21, 31, 4 1 were obtained by transferring the original parents to bottles 100001, 200001, 300001, 400001 respectively - as described in the seotion on heat treatment. These controls were set up on January 31 at 5:00 p.m. and parents were removed on February 3 at 5:00 p.m. The Controls were kept at a temperature of 25° C. P^  emerged on February 10 - results are listed in Table XXXII. Table XXXII flOOOO1 'Hot mal 9? Synj o V >alpi 99 fgoooo 1 Nor mal 99 Synp alpi 99 Feb. 12 14 Total 40 11 48 13 Feb. 12 14 Total 23 36 37 32 51 61 0 0 59 69 0 0 #300001 Feb. 12 14 Total 13 0 10 0 #400001 Feb. 12 14 Total 65 21 67 24 13 10 0 0 86 91 0 0 F 1 interses of controls were set up: B #100001 - 3 P 1?? x 3 F/W- B #A - February 12 200001 ' •» 3F 1 99 X 3 I^cPcFL B #B - February 12 300001 - 3 F 1?? x 3 f ^ o V - B #C - February 12 400001 - 3 . F l ^ x 3 . ? l o V , - B f D - February 12 F 2 results of oontrols are listed in Table XXXIII. - 36 .-Table XXXIII A normal Synpalpi B normal Synpalpi o-V ?9 cy<y 9% 99 9? Feb.24 0 0 Feb.24 2 0 1 25 4 5 1 0 25 9 4 2? 31 30 27 10 12 28 23 17 28 11 11 Total 58 52 1 0 Total 32 30 0 l C D Feb.24 59 52 0 1 Feb*24 16 22 1 l 25 19 24 25 14 12 0 i 27 39 26 27 54 60 0 2 28 11 11 28 4 5 Total 128 113 i 0 1 Total 88 99 1 4 As a control for the whole oold treatment, orosses were set up using wild stook in order to test the lethality at 15° Oi On February 24 at 5 p.m. a cross was set up: #1 12 9$ x 120-Vali wild type flies (Urbana S stock) On February 25 at 5 p.m. the parents were transferred to #10. On February 26 at 5 p.m» the parents were transferred to #100. On February 27 at 5 p.m. the parents were transferred to #1000. On February 28 at 5 p.m. the parents were removed from #1000. On February 28 at 5 p.m» B #1 was placed in the cold (15° C.)-age of culture was 3-4 days. On March 3 at 5 p.m. B #10 and #100 were placed in the cold. Age of #10 was 5-6 days, and #100 was 4-5 days. B #1000 was used as a control and was left at 25° C. Bottle #1 was removed from the oold on March 14 - and #10 and #100 were removed March 17. 37 Total F 1 counts of ?9 wild oontrols: cry? 81 f l 77 #10 85 80 #100 87 81 #1000 90 93 F* interses were set up and F 2 total counts were: #11 120 125 #101 134 130 #1001 127 125 #10001 125 121 The results of the preceding cold treatments have been sum-marized and tabulated in Table XXXIV. Table XXXIV - Cold Treatments Cross NO. Consti-tution. Age When Treated No. of pi Emerged F l Syn-palpi Life History No. of F 2 P 2 Syn-palpi Dissection of F-*- Pupae 1 1 13 Xple?? x 15 Xpleo*^ 4-5 days 2 .0 Dec.18-Jan.20 In cold Dec.23 Out Jan.13 0 0 NO larvae Sampling of 22 pupae Emergent: 1800*3?$ Larval : 1 10 13 Xple?? x 15 Xple 6* a* 3-4 days 70 36^"34?? 0 Dec.19-Jan .25 In cold Dec.23 Out Jan. 13 145 66°-V79 ?? 0 No larvae 19 pupae Emergent: 13 (3 syn) Larval : 1 100 13 Xple?? x l 5 Xple 6*0" Control (25° C ) 134 10 {8%) Dec.20-Jan. 2 Interse: 114 5 4 0 V 6 O ? £ * P 1 syn. Interse: 122 5 8 0 V 6 4 ? ? 4??(4#) 10 (Q%) 50*0* 5 ?? 2 14 Xple?? x l 5 XpleoV 4-5 days 0 0 Dec.18 In cold Dec.23 Out Jan.13 0 0 No larvae 44 pupae Emergent: 340^3?? Larval : 7 Sinpa\p\ Table XXXIV (Continued) Cross No. Consti-tution Age When Treated No* of F Emerged p l Syn-palpi Life History No. of F* F^ Syn-palpi Dissection of P 1 Pupae 20 14 Xple ¥ ¥ x l 5 Xplecf^ 3-4 days 90 45°-V45?? 0 Dec.19-Jan.25 In cold Dec.23 Out Jan.13 162 6 9 0 V 92?? 1 V No larvae 28 pupae Emergent: ,jqc?b»30$ Larval : 7 200 14 Xple 2¥ x l 5 Xple^o* Control (25° c) 132 65<yV67?? 3 (2%) 2oVl % Dec.20-Jan. 2 Interse: 147 69«V72 ??• P 1 syn. Interse: 124 SOoV6 4 99 692(3%) 7 ( # ) 3 0 V 4 $ ? 2 1 12 Xple?? x28 Xple Co* 3-4 days 178 2??(#) Jan.27-Feb.17 In cold Jan.31 Out Feb.14 Interse: 73 3 8 ^ 35 n F 1 syn.? x F 1 0^ 59 3 6 0 * 0 * 2 5 ? ? 0 0 No larvae No pupae 2000 12 Xple** x 28 Xple o" a" 5 - 6 days 0 0 Jan.24 In cold Jan.30 Out Feb.12 0 0 No larvae Many pupae -sample of 15 Emergent: 5oV7^ Larval : 3 200003 12 Xple ?? x28 Xplec^o71 Control (25° C) 128 0 Jan.31-Feb.10( 63 32 0 ^ 3 1 ? ? 19(2%) Table XXXIV (Continued) Cross No© Consti-tution. Age When Treated No. of F Emerged p i Syn-palpi Life History No. of F 2 P2 Syn-palpi Dissection of F 1 Pupae 3 1 Xple Synpalpi? 3 Xple Synpalpi 4-5 oays No eggs < >r larvae 30 tt 3-4 days ti I I I I it. 300 n Control (2SO c) I I I I it I I i 1 3 Xple ?? Synpalpi 1 Xple Synpalpi 3-4 days 26 13.0-V13?? 0 Jan.27-Feb.17 In cold Jan .31 Out Feb.14 53 0 No larvae 15 pupae Emergent: 5°V10?? Larval : 0 1000 5-6 days 0 0 Jan .24 In cold Jan.30 Out Feb.12 0 P No larvae Many pupae -sample of 15 Emergent: 9 o V 6 ? S Larval : 0 100001 lit Control (25° O 112 51cV6l¥¥ 0 Jan.31-Feb.10 111 5 5 Xple orange ?-$ KlO Xple orange © V 4-5 days No eggs c r larvae 50 tt 3-4 days 0 0 Dec.19-In cold Dec.23 0 0 No larvae 24 pupae Emergent: l l o V 3 ? 2 Larval : 10 Table XXXIV (Continued) Cross No. Consti-tution. Age When Treated No. of p l Emerged P l Syn-palpi Life History No. of P 2 P 2 Synpalpi Dissection of P 1 Pupae 500 5 Xple orange 99 x 10 Xple orange o-V1 Control (25° C) 51 25oV26?? 9 (18$ 4 0 V 5 ?$• Dec.20-Jan. 2 Interse 130 60oV709? P 1 Syn. Interse 114 53crv61?? 12 (9%) lcr> 119$ 8 (7%) 3<3*6» 5?$ 6 5 Xple orange 99 x 11 Xple orange <rV 4-5 days 0 0 Dec.18-Jan.20 In cold Dec.23 Out Jan.13 0 0 No larvae 30 pupae Emergent: 21oMl syn) Larval : 9 60 ti 3-4 days 0 0 Dec.-19 In cold Dec.23 Out Jan.13 0 0 No larvae 31 pupae Emergent: l l c V l ? Larval : 19 600 tt Control (25° C) 74 290-V-45?? 12 (ie$) 2oV»10?? Dec.20-Jan. 2 Interse 121 560V65?? P 1 Syn. Interse 118 59°-V59?? 9 (7.30 10 {&fo ) 40^6 ?¥ Table XXXIV (Continued) Cross No. Consti-tution Age When Treated No. of p i Emerged p i Syn-palpi Life History No. of F 2 Syn-palpi Dissection of F-*- Pupae 8 Xple orange ?? x 10 Xple orange cr'cri 3-4 days 1 4 « 13 o^i?^ 0 Jan.27-Feb.17 In cold Jan.31 Out Feb.14 " 0 0 No larvae 94 pupae Emergent: 61^0^30?$ Larval : 3 4000 i t 5-6 days 0 0 Jan.24-In cold Jan.30 Out Feb.12 0 0 No larvae Many pupae - sample of 15 Emergent: l O o-ic 1 3?S Larval : 2 40000 it . Control (25° C) 177 860^91?$ 0 Jan.31— Feb.10 192 89o^l03??-5 (2.90 4?? 7 2; Xple orange syn. ?9 x 4 Xple orange syn. cr^cyi 4-5 days. 0 0 Dec.18-In cold Dec.23 Out Jan.13 0 0 No larvae 34 pupae Emergent: l l c ^ l syn) 499 Larval : 19 ,70 It! 3-4 days 0 0 Dec.19-In cold Dec. 23 Out Jan.13 0 0 No larvae 41 pupae Emergent: 120*0(7 syn) 3 9S Larval : 26 Table XXXIV (Continued) >oss No. Consti-tution. Age When Treated No* of p i Emerged p l Syn-palpi Life History No. of F 2 F* Syn-palpi Dissection of F pupae 700 2 Xple orange syn. ??• x 4 Xple orange syn. o-~V Control (25° C) 57 21o^36?? 24 ( 4 # ) 7o->-a799 Dec.20-Jan. 2 Interse 120 560V64??. F 1 syn. Interse 123 59o-v6"4?$ p l syn? X P"*- Cf) 52o^52 9? is m>) 7oV»8 9% 15 (125?) 6c^9 9$ 3p-">(ljO 2 Xple orange syn. ¥9 x 3 Xple orange syn. cr^o77 3-4 days 2 o ' V 0 Jan .27-Peb.17 In cold Jan.31 Out Feb.14 0 0 No larvae 2 pupae Emergent: 2 ? $ Larval : 0 3000 ti 5-6 days 0 0 Jan .24-In cold Jan.30 Out Feb.12 0 0 No larvae Many pupae-sample of 15 Emergent: 4 O 0 V 8 9 $ Larval : 3 30000 tt Control (25° c) 23 13o>b*L0?? 0 Jan.31-Feb.10 242 12 803114?? 19- (.50) 44 Cold Treatments Observations: In all oold experiments the age groups of 4 - 5 and 5-6 days proved most critical when subjecting the Xple, Xple synpalpi, Xple orange, and Xple orange synpalpi cultures to a cold treatment of 15° C. Of the two groups the 5-6 day series was the more sensitive - no P1 emerging in any of the crosses. Very few F* emerged in the 4-5 day series. The cold reduced the number of synpalpi appearing in the F 1 and F 2 - in all oases the controls produced a fair percentage of synpalpi (from 7$ to 12$). - 45 -(it) Heat treatments* Temperatures of 35° C* and 29° 0. were used in heat treatments* On January 26 at 5 p.m* the following crosses were set up: #101 - 3 Xple synpalpi ¥?x 1 Xple synpalpi o? #201 - 12Xple¥? x 28 Xpleo^ (from Xple synpalpi stook) I301 - 2 Xple orange synpalpi??x 3 Xple orange synpalpio-V #40* - 8 Xple orange?? x 10 Xple orange ^ V?( from Xple orange synpalpi stock) On January 29 at 5 p.m. the parents were transferred to new bottlest #101 to #1001* #201 to #2001, #301 to fSOO1, #401 to I4001. On January 30 at 5 p*m. the parents were transferred again to new bottles: #1001 to #10001, #2001 to #20001, #3001 to #30001, #4001 to #40001* On January 31 at 5 p*m. the parents were transferred to new bottles: flOOO1 to flOOOO1, §Z000l to #20000*, #3000* to #30000*, #4000* to #400001* On February 3 at 5 p.m* the parents were removed from bottles #10000*, #20000*, #30000*, and #40000** These were used as controls. On February 3 at 11:15 a.m. the bottles of series #10* 20* etc. (age, 5-6 days), #1001, 2001 etc. (age, 4-5 days), and #10001, 2000* etc* (age, 3-4 days) were placed in the heat at 35° C. These bottles were removed from the heat on February 10 at 10 a.m. No F* flies emerged in any of the heat-treated cultures - a l l died in pupal or larval stages. The bottles were examined and the pupa oases were opened. Heisults of this procedure are listed below* - 46 -B #10 Ho larvae Sample of 12 pupae B #20* 3 larvae Sample of 12 pupae 1 B #30 No larvae Sample of 12 pupae B #40 No larvae Sample of 12 pupae B JlOO1 A few larvae (dead) Pupae B #2001 Some larvae (dead) Pupae B #300* No larvae or pupae visible. B fiOO1 No larvae Pupae B #1000* A few larvae (dead) B #2000* A few larvae (dead) B #30001 A few larvae (dead) B #4000* Many larvae (dead) Emergent: Larval : 8o-V4?$ 0 : Emergent: 8^3?? Larval : 1 : Emergent: Larval : Emergent: Larval : 0 2 cV"? 10 : Emergent; 0 Larval : all : Emergent: 0 Larval : all : Emergent: 0 Larval : a l l no pupae no pupae no pupae no pupae - 47 --The controls #10000*, #20000*, #30000*, #40000*, were kept at a temperature of 25° C. The F* results are listed in Table XXXII and the F 2 results, in Table XXXIII, On March 11 at 5 p,m# the heat experiment was repeated exactly as the previous one except that a temperature of 29 C. was used instead of 35° C, In place of the numbers 10 1 - 20* eto, numbers 10*, 20j£ etc, were used, in place of numbers 100* etc, numbers 100^ etc,, in place of numbers 1000* etc, numbers lQOoj" etc., and in place of numbers #1000* etc, ( 3 - 4 days) were placed in 29° C, chamber on March 17, 5 p,m, and removed on March 24 at 5 p*m. Bottles #10000." etc. were used as intersed and produced 72cfVand 76"?? - no synpalpi appearing. All heat treated bottles were examined. The observations are noted below: B #10* No larvae 108 pupa cases : Sample of 15 Emergent: 6o-V74?$ Larval : 5 No larvae 120 pupa cases : Sample of 15 Emergent : ¥fWb<& Larval : 6 - 48 #30* No larvae A 101 pupa oases : Sample of 15 Emergent : 6 trV4,9$ larval : 5 No larvae 115 pupa oases : Sample of 15 Emergent : 9 0*6* 3 Larval t 3 10 flies partly emerged #100^  No larvae 105 pupa cases : Sample of 15 Emergent : 4 cPa* 4 9% Larval : 7 #200^  No larvae 112 pupa cases t Sample of 15 Emergent ; 4 6*0-1 3 £5 Larval : 8 #300?" No larvae 107 pupa oases : Sample of 15 Emergent : 6 o-V 2 9-S Larval : 7 No larvae 108 pupa cases : Sample of 15 Emergent s 6 o^a77 3 99 Larval : 6 #1000A No larvae 109 pupa cases t Sample of 15 Emergent: 4 0 ^ 0 ^ 1 ? Larval : 10 #2000^  No larvae 119 pupa cases : Sample of 15 Emergent : 2 o-V"7 Z99-Larval : 11 - 49 #3000' A No larvae 103 pupa oases Emergent Larval #4000^  No larvae 126 pupa oases Emergent Larval Sample of 15 3 ofo-i 1 $ 11 : Sample of 15 i 3 crlcP 2 $ t 10 P 1 results of controls are listed in Table XXXV along with the 2 2 1 F counts. F counts were the result of F interses. Table XXXV #10000* Non o-">cr> sal ?? Synpj ilpi #20000| Non c r V mal 99 Synp alpi 99 pi •g2 63 50 67 51 1 pi F 2 71 37 75 48 3 #30000^  F 1 F 2 69 66 72 63 1 #40000j F 1 F 2 63 67 82 68 1 2 As a control for a l l heat treatments, wild cultures of the three ages were subjected to similar temperatures as those used in Xple orosses - the lethality of these temperatures on ordinary wild stock being observed. On February 24 at 5 p.m. the following crosses were set up using Urbana-S wild stock: #1 12oVfe 12 9$ #2 12CVJC 12 ?? On February 25 at 5 p.m. the parents were transferred to new bottles: #1 to #10, #2 to #20. -50 On February 26 at 5 p.m. another transfer of parents was carried out: #10 to #100, #20 to #200. On February 27 at 5 p.m, a f i n a l transfer of parents occurred: #100 to #1000, #200 to #2000. On February 28 at 5 p»m. the parents were removed from #1000 and #2000, At 5 p,m, on February 28 B #1 (age 3-4 days) was placed in heat at 35° C. and B #2 (age 3-4 days) i n heat at 29° C. On Maroh 3 at 5 p.m. B #10 (age 5-6 days) and #100 (4-5 days) were placed i n heat at o 35 0 , and B #20 (age 5-6 days) and #200 (4-5 days) were placed i n heat at 29° C. B #1000 and #2000 were used as.controls and were kept at the ordinary temperature of 25° C. F and F counts are l i s t e d i n Table XXXVI - the F counts being the result of interses. Table XXXVI #1 ' ?? #3 .0 ?$ #K 30 #1 000 «« £0 ?¥ £ £00 9 ? #2C 100 92 F 1 F 2 67 112 64 118, 72 121; 68 118; 74 98 69 1P7 83 106 78104 83 124 70 127 78 117 70 123 80 105 73 111 89 130 85 135 A l l f l i e s were normal wild type - the heat treatments had no to effect other than^speed up l i f e histories. No f l i e s died i n larval or pupal stages. A l l the results from heat treatments are summarized and tabu-lated i n Table XXXVI.I. Observations: The Xple stocks were very sensitive to heat - temperatures of 29° and 35° C. k i l l e d a l l - so that no F 1 emerged. The percentage of synpalpi in the controls was lower than usual. Table XXXVII - Heat Treatments Cross No. Consti-tution. Age When Treated Temp. C. No., of F Rmereed pi Syn-palpi Life History No. of p2 T2 Syn-palpi Dissection of F pupae IO*- 3 Xple syn. 1 Xple syn. 5 -6 days 3 5 ° 0 0 Jan.28-In heat Feb. 3 Out Feb.10 0 0 No larvae Sample of 12 pupae Emergent: 8 c / V 4 ? $ Larval : 0 1 0 0 1 11 4-5 days 3 5 * 0 0 Jan.29-In heat Feb. 3 Out Feb.10 0 0 A few dead lapvae Sample of 12 pupae Emergent: 0 Larval : 12 1 0 0 0 1 3 it 3 - 4 days 3 5 ° 0 0 Jan.30-In heat Feb. 3 Out 0 0 A few dead larvae Pupae - 0 lococr it Control 25° 112 51<^61? 0 Jan.JL -Feb.10 111 5 9 0 ^ 2 $ l°t'9#) tt 5 - 6 days 290 0 0 Mar«10 In heat Mar.17 Out Mar.24 0 e No larvae 108 pupae - sample of 15 Emergent: 6 ^ 4 ? $ Larval : 5 1 0 0 i ti 4 - 5 days 2 9 ° 0 0 Mar.12-In heat Mar.17 Out Mari24 0 0 No larvae 105 pupae - sample of 15 Emergent: 4o-V4 9$ Larval : 7 1000} A tt 3 - 4 days 2 9 ° 0 0 Mar .13-, In Mar. 17 Out Mar.24 0 0 No larvae 109 pupae -sample of 15 Emergent: AcrV'l ? Larval : 10 Table XXXVII - Heat Treatments (Continued) 3ross No© Consti-tution Age When Treated Temp. C. No.-.of P Emerged ^Syn-palpi Life History No«9of P p2 Syn-palpi Dissection of P pupae toooo^ 3 Xple syn.?? 1 Xple syn.o'7 Control 25° 130 630*67?? 0 Mar«,14«* Mar.24 101 50o-^ 51?? 1^(1$) 2 0 1 12 Xple 28 Xple 0*0* 5-6 dajs 35° 0 0 Jan.28«» In heat Feb. 3 Out Feb.10 0 0 3 larvae Sample of 12 pupae Emergent: QorW3 ? 9 Larval : 1 2 0 0 1 in 4-5 days 35° 0 0 Jan.29-In heat Feb. 3 Out Feb.10 0 0 Some larvae Pupae: -Emergent: 0 Larval : A l l 2000 1 11 3-4 days 35° 0 0 Jan.30-In heat - Feb . 3 Out Feb.10 0 0 A few dead larvae Pupae: 0 20000 1 it Control 25° 128 59069$ 0 Jan.31-Feb.10 63 32051? 1 ? ( # ) 20i it 5-6 day3 29° 0 0 Maro i l -In heat Mar.17 Out Mar.24 0 0 No larvae 120 pupae - sample of 15 Emergent: ^cr'cy'^^ Larval : 6 200* it 4-5 days 29° 0 0 Mar.12-In heat Mar.17 Out Man>^  0 0 No larvae 112 pupae-sample of 15 Emergent: 4 0 ^ 3 ? $ Larval : 8 Table XXXVII - Heat Treatments (Continued) Cross No, Consti-tution. Age When Treated Temp.' C. No.-.of F Emerged p i Syn-palpi Life History No. of F^ F 2 Syn-palpi Dissection of F pupae 2000* 12 Xple?? X 28 Xple o-V 3-4 days 29° 0 0) Mar.13-In heat Mar.17 Out Mar.24 0 0 No larvae 119 pupae - sample of 15 Emergent: 2<rV2 9? Larval : 11 20000* ti Control 25° 146 0 Mar.14-Mar.24 75 37°'"748¥ 3 ? <A%) 30 1 2 Xple orange syn« ?? X 3 Xple orange syn 0 o*cri 5-6 days 35° 0 0 Jan.28~ In heat Feb. 3 Out Feb.10 0 0 No larvae Sample of 12 pupae Erne rge nt: 9o-V 3 9 9 Larval : 0 300 1 it 4-5 days 35° 0 0 Jan.29-In heat Feb. 3 Out Feb.10 0 0 0 3000 1 It; 3—4 days 35° 0 o. Jan.30-In heat Feb. 3 Out Feb.10 0 0 A few dead larvae Table XXXVII - Heat Treatments (Continued) Cross No. i Consti-tution. Age When Treated Temp. C. No. of F x Emerged Syn-palpi Life History No. of p2 P 2 Sym-palpi Dissection of F pupae 50000x 2 Xple orange syn. X 3 Xple orange syn. o^o^ Control 2 5 ° 23 13cU0? 0 Jan.30-Peb .10 242 L28 o"0'7 1 1 4 ? ? 1?(.S*) i n 5 - 6 days 29° 0 0 Mar.11-In heat Mar.17 Out Mar.24 0 0 No larvae 101 pupae - sample of 15 Emergent: 6 0 V 4 ? 0 . Larval : 5 5 0 0 J i t 4-5 days 29° 0 0 Mar.12-In heat Mar.17 Out Mar.24 0 0 No larvae 107 pupae - sample of 15 Emergent: 6 o^cr>2 ?$ Larval : 7 300C-1 It 3-4 days 29° 0 0 Mar.13-In heat Mar.17 Out Mar.24 0 0 No larvae 103 pupae - sample of 15 Eme r ge nt: 3 o ^ l ? Larval : 11 50000* t » Control 2 5 ° 141 6 9 0 = 7 2 ? 0 Mar.14-Mar.24 131 68o-">o-"> 6 3 ? ? 1S(.9*) Table XXXVII - Heat Treatments (Continued) Cross No. Consti-tution. Age When Treated Temp. C. No...of F X Emerged P 1 Syn-palpi Life History No. of F F 2 Syn-palpi Dissection of F pupae 8 Xple orange ?? X 10 Xple orange cr'0'7 5-6 days 35° 0 0 Jan 028-In heat Feb. 3 Out Feb.10 0 0 No larvae Sample of 12 pupae Erne rge nt: 2 c r 7 cr* Larval : 10 400 1 it 4-5 days 35° 0 0 Jah.29* In heat Feb. 3 Out Feb.10 0 0 No larvae Many pupae Emergent: 0 Larval : A l l 4 OOO1 tr 3-4 days 35° 0 0 Jan.30** In heat Feb. 3 Out Feb.10 0 0 Many larvae No pupae 400001 tt Control 25° 177 s e ^ i ? 0 Jan .31"1 Feb.10 192 89<y7<f 103 5 (2„5#) l o * 4?$ 40*. tt 5-6 days 29° 5 3«^2$ 0 Mar.11-In heat Mar.17 Out Mar.24 148 76 n 0 No larvae 115 pupae - Sample of 15 Erne rge nt: 9 o-V>3 Larval : 3 10 partly emerged. Table XXXVII - Heat Treatments (Continued) Cross; No. Consti-tution. Age When Treated Temp. C. No. of p i Emerged P 1 Syn-palpi Life History No. of p2 p2 Syn-palpi Dissection of pl- phpae 400* 8 Xple orange 9? 10 Xple orange <rV 4-5 days 29° 0 0 Mar.12-In heat Mar.17 Out Mar.24 0 0 No larvae 108 pupae - sanple of 15 Emergent: 6 0 ^ 3 9 0 Larval : 6 40001 »t, 3-4 days 29° 0 0 Mar.13-In heat Mar.17 Out Mar.24 0 0 No larvae 126 pupae - sample of 15 Emergent: 3crkP299 Larval : 10 400001 A it Control 25° 165 830*829-0 Mar .14** Mar.24 135 67 Co* 6899 3 (2%) lo* 299 - 5 7 -( i i i ) Discussion; In conjunction with both heat and cold treatments of Xple stock, wild stocks were also subjected to similar temperatures. A l l f l i e s emerging from these treatments were normal - the F 1 and F 2 counts being normal. This indicates that the Xple stock i s extremely sensitive to variations from the normal temperature of 25° C - since 15° C. re-duced the number of synpalpi appearing and was lethal in some age groups, and since 29° C. and 35° C. were both lethal temperatures for the Xple stock. (d) Investigation of the lethality of the mutant The possibility of a lethal character being associated with the mutant, synpalpi, was investigated in two ways. The f i r s t method was to examine cultures of Xple and Xple synpalpi for lethality before emergence and the second was to test with C1B stock. This investigation was carried out since the preliminary crosses suggested a lethality and since Zaliss (1939) described a deficiency affecting the X chromosome whioh caused lethality in the young larvae and eggs. (1) Crosses of Xple and Xple synpalpi. On Apr i l 1 at 5 p.m. the following crosses were set up: A - 4 Xple synpalpi?? x 3 Xple synpalpi o^c/i B - 4 Xplfl ? ? x 3 Xple o"o» C - 4 Xple orange synpalpi?? x 3 Xple orange synpalpi <rV D - 4 Xple orange?? x 3 Xple orange^c" 1 On Apr i l 2 at 5 p.m. the parents were transferred to new bottles: l 1 1 1 A to A , B to B , C to C , and D to II . On A p r i l 3 at 5 p.m. another transfer of parents was made to new bottles: A 1 to A 2, B 1 to B 2, C 1 to 0 2 and D 1 to D2. o 2 2 2 The parents were removed from A , B , 0 and D on A p r i l 8» An account of the date of emergence, the number of f l i e s that emerged, and the number that died before emergence i s given below: A: Xple syn? x (emerged Apr i l 10: 21cf>32? - no synpalpi Xple syno* (died in pupa case: 4 - 1? , 1°^ , 2 larval (^  .dead : 6% - 59 A : Xple syn? x Xple syno^ A : Xple syn? x Xple syn 0-7 B : Xple? x Xple °^  BA : Xple? x Xple B ; Xple? x Xpleo-7 C t Xple orange syn. ? x Xple orange syn • cr> C* : Xple orange syn ? x Xple orange syn cr? G2 : Xple orange syn.? x Xple orange syn. 0* D : Xple orange ?x Xple orange D : Xple orange?x Xple orange °^  D s Xple orange?x Xple orange emerged Apr i l 11 : 29 o^M? -died in pupa case: 2 - l f f l , $ dead : 5$ control: emerged Ap r i l -12 : 48 0^50$ -emerged April 13 : 66o^ 63? -died in pupa case: 3-1 Or* 1? , fa dead • : 2.3$ fa emerged A p r i l 12 : 68 CT1 66$ -died in pupa case: 3-2 o» 1? $ dead : 3$ control: emerged Ap r i l 12 : 430* 39 ¥ -emerged A p r i l 10 : 48 o71 48? -died in pupa case: 2 - ley, 1? $ dead : 2$ no synpalpi 2?, 1 or> 1 larval fa synpalpi: 2.3$ emerged Ap r i l 11 died in pupa case $ dead control: emerged Ap r i l 12 : emerged Ap r i l 11 : died in pupa case: fa dead : emerged Apr i l 11 : died in pupa case: fa dead : control: emerged Apr i l 12 : 240" 43? - no synpalpi 2 3 ^ 35? - no synpalpi 52 54? - synpalpi: 3? 16: 8or> 4?) 4 larval 15$ $ synpalpi: 2$ 470-T 63? - synpalpi: 1? 8: l < r » 4?, 3 larval 7$ fa synpalpi: .9$ 510^ 57? - no synpalpi 60 -In order to determine the number of wild type f l i e s that died before emergence 2 crosses v/ere set- up and examined - thus establishing a control for the above Xple orosses. * B #1 : ZcrVx 4??- B #2 : 4o-Vx 4 ? ? The parents were transferred to new bottles #10 and #20 respectively at the end of 24 hours. The parents were removed from B #10 and #20 at the end of the second'24 hours, thus setting a standard time for deposition of eggs. A l l f l i e s emerged in 9 days* time - an account of the number emerged and the number that died before emergence i s given below: B #1 number emerged: 740^85?-number dead : 0 B #10 number emerged: 89 cr> 91 ? number dead : 0 B #2 number emerged: 72 73? number dead : 0 B #20 number emerged: 104 102? number dead : l&i.Bfo) Observations: Since no wild f l i e s died before emergence and since 2 - 15$ the Xple and Xple orange f l i e s died before emergence, i t may be assumed that there i s some factor br factors)present in these Xple f l i e s causing their death or reduced v i a b i l i t y . The percentage of death and / synpalpi varied considerably but i t was significant that the Xple and Xple orange crosses produced 2-3$ synpalpi 'whereas the Xple synpalpi N- 61 -(whereas the Xplo synpalpi) and Xple orange synpalpi crosses produced none. Death before emergence was highest in the Xple orange cross (average 11.5$) and in the Xple synpalpi cross (average 6.5$). In the dissection of the pupa cases i t was again noted that several of the f l i e s were attached to the end of the pupa cases by means of the tracheae, thus being unable to work themselves free. In their efforts to free themselves, they contracted and expanded their bodies, forcing the front of the head out i n the region of the antennae. As mentioned before this forcing of the head out may weaken i t and force the antennae together - since the front of the head of synpalpi f l i e s i s bulged. This attachment of the f l i e s to the end of the pupa case and subsequent elongation efforts may account for the squeezed appearance of the abdomen of some of the synpalpi f l i e s which managed to extricate themselves. (2) C1B method. The C1B method of detection of lethal mutations i3 limited in that only sex chromosomes are involved i n the test. The GIB female used in the test has on one of i t s X chromosomes three factors on which this method is baseds a known lethal gene, 1, which k i l l s the males that receive this chromosome, sinoe these receive no normal a l l e l e in the - 62 -Y ohromosome; a factor C, which prevents crossing over in the X, thus insuring that this chromosome will not lose its lethal; and a dominant gene, B (Bar eye) which marks this chromosome - thus making it possible to identify the offspring that receive i t . The prooedure is to cross a male containing the mutation with a C1B female. Prom this cross the Bar female offspring (which contain the C1B chromosome and the.mutation chromosome) are crossed with the normal type F 1 males. Such a cross produces Bar and not-Bar females, half the males die because of the C1B lethal, and if the mutation tested is a lethal the other half of the males will also die. Visible mutations may be detected by the examination of the surviving F 2 males. On February 21 the following crosses were set up: (all females used were virgin). A 1 ClB/dl 49 ? x 1 Xple orange synpalpi cr1 B 1 ClB/dl 49? x 1 Xple orange °^ C 1 lz/ClB x 1 Xple orange synpalpi 0-^  D 1 lz/ClB ¥ x 1 Xple orange ^ E 1 lz/ClB x 1 Xple synpalpi cr> F 1 lz/ClB x 1 Xple o"1 0 1 lz/ClB 9. x 1 Xple synpalpi cr7 H 1 lz/CiB ? x 1 Xple & I Control: 1 lz/ClB ?- x 1 wildo-7 The F A counts are listed in Table XXXVIII - 6 3 -Table XXXYIII A B C D E P 0 H I C1B ?¥ 45 26 35 36 14 41 26 19 26 H ?? 28 21 40 32 15 41 29 18 25 H (TV 29 30 35 28 6 34 17 16 28 F interses were set up: 1 F 01B? x 1 F cr"7 2 F counts are listed in Table XXXIX. Table XXXIX Bottle CIS?? B?? Synpalpi F* orcf} % Syn. A 128 134 120 5 1.3 B 138 132 114 12 3 C 109 125 131 4 1.1 D 123 148 126 5 1.2 E 141 140 140 0 0 F 124 128 115 9 2.4 a 148 148 145 0 0 H 86 82 58 0 0 I 133 134 126 0 0 In the above table N ¥- and No-7 refer to those flies whioh re-ceived the homologous X chromosome (not C1B) from the C1B mother. Ho lethal appears to be associated with the vfirst chromosome sinoe the ratios were as expected: 1 C1B?- : 1 N? : l o 7 , Synpalpi did appear in some of the males - this seems to indicate that it is associated with the X chromosome in some way. (iii) Disoussion: This investigation of the lethality of the mutant has shown that there is some faotor whioh causes the death of the Xple, Xple synpalpi, Xple orange, and Xple orange synpalpi flies before emergence, The C1B method of testing showed that there is no lethal factor as-sociated with Chromosome I alone. This method is extremely limited since it tests only Chromosome I; the lethality, therefore, may he as-sociated with Chromosome II, III* or IT alone or a combination of any of the Chromosomes. Subsequent tests for linkage group may reveal the association of this lethal factor. - 65 -5* Determination of the linkage group* In order to determine the linkage group with whioh the mutant synpalpi is associated standard testing stocks were used* (I) Ohromosome I In the test for linkage with Chromosome I yellow stock was used - yellow being located on Chromosome I. On February 14 the following cross was set up: 1 yellow synpalpi? x 2 Xple orange synpalpioV, F 1 oounts are listed in Table XL. Table XL Ye! Llow Wild Hon o-V ml ?? Synp o V alpi 99 Hon crV nal Synpj ilpi ?? Feb.24 25 26 2? 28 Mar. 3 Total 24 13 19 13 13 4 15 7 7 4 1 6 5 8 5 3 4 5 66 40 0 0 0 30 0 6 F crosses were set up on February 25. 3 F 1 yellowest 3 F 1 yellow o-V 3 F virgin wild?* x 1 Xple orange synpalpi C7 3 F virgin yellow?? x 1 Xple orange synpalpicr> 3 F 1 virgin wild¥?-x 2 F 1 yellow aV7. The following 1 Y : 2 Y : Y 3: Y 4: The F 8 results are listed in Table XII. -66-Table XLI Yellow . Kor o-V mal 99 Synp alpi 9$ Nor mal 99 Syni jalpi 9 ? Y1 Mar.12 14 Total 53 25 40 36 78 76 0 0 0 0 0 0 Y 2 Mar*12 14 Total 22 6 16 6 41 10 2 3 3 26 0 0 0 22 51 2 6 Y 3 Mar*12 14 Total 41 27 46 28 68 0 0 0 0 74 0 0 Y* Mar*12 14 Total 28 10 12 12 10 16 23 12 2 0 36 24 0 0 26 35 2 0 Expected resultst t Y1 - a oross of F 1 yellow ? x F 1 yellow cr7- since yellow appears only in a homozygous oondition: the F 2 results should he: yellow or1 and yellow^ in a 1:1 ratio* Y2 - a cross of F 1 wild? x Xple orange synpalpi cf1 - since the F 1 wild? will have one homologue from the yellow parent and the other from the Xple synpalpi parent, the F 2 results should he: 2 Xple9 t 1 Xpletf"1: 1 yellower7. Y3 - a cross of F 1 yellow? x Xple orange synpalpi cr7 - since the F* yellow ¥ is homozygous for yellow, the F^  results should he: 1 Xple 9 i l yellow cr7. Y^  - a oross of F 1 wild? x F 1 yellowo^- sinoe the F 1 wild$ will have one homologue from the yellow parent and the other from the Xple synpalpi parent, the F 2 results should he: 1 Xple* :1 yellow? :1 Xple©*: 1 yellower^ 67 * It was believed that the yellow synpalpi female had been pre-fertilized by a yellow male, sinee the cross of yellow?, x Xple orange cr7 gives only yellow males and Xple females, whereas the actual F 1 count included yellow males and females as well as wild females. This assump-tion was further substantiated by the F 2 results. The actual F 2 results conformed to the above expected ratios. 2 4 Synpalpi flies appeared in crosses Y and Y in Xple flies whioh were homozygous for the Xple synpalpi horaologue obtained from the original Xple synpalpi male parent. In cross Y 2 the F 2 oonsisted of Xple males and females and yellow males. Half of the Xple females were homozygous for the Xple synpalpi homologue obtained from the original Xple synpalpi male parent and half were heterozygous for it, having one yellow X homologue and one Xple synpalpi X homologue. The yellow males had the one X homologue derived from the yellow female parent and the Xple males had the one X ohromosome derived from the Xple synpalpi male parent. Synpalpi ooourred in some Xple females and in some Xple males, presumably in those whioh were homozygous for synpalpi. In cross Y4 the F 2 con-sisted of 2 types: yellow males and females, and Xple males and females -the Xple females were heterozygous for Xple synpalpi and yellow, therefore only in the Xple males was the Xple synpalpi homologue homozygous -Synpalpi appeared only in Xple males in this cross. In no other crosses did the Xple synpalpi X ohromosome from the original synpalpi male parent appear doubly in the female or singly in the male - thus making it homozygous - and in no other crosses did synpalpi appear. This seems to indicate that synpalpi is associated with the X chromosome, appearing when it is homozygous. «• 58 *• (ii) Chromosomes II and III* In the test for linkage with Chromosomes II and III, Star > Diohaete stock was used - S/Cy D3/ln (3L) PMe - Star located on Chromosome II, Dichaete on Chromosome III. On February 14 the following cross was set up 2 Star Dichaete virgin ??x 2 Xple orange synpalpi 0 ^ On March 3 a cross was set up of F 1 virgin curly dichaete? x F 1 curly diohaete cr7 , The F 2 results are listed in Table XLII. Table XLII Curly dichaete Curly Dichaete ? Xple &> a* ?? cV n o-V 9% Mar*14 22 26 6 9 11 9 6 7 17 20 32 19 9 13 14 3 9 18 5 8 7 4 10 5 1 3 19 8 9 4 5 2 1 2 2 Total 55 75 36 27 36 29 12 21 ? Xple refers to the fly with both homologues of the 2nd and 3rd chromosomes obtained from the mutant* The expected results were: 4 Curly Dichaete: 2 Curly: 2 Dichaete: 1 ? Xple* The actual F2 counts followed this ratio very closely: 130 Curly Dichaete: 63 Curly: 55 Dichaete: 33 ? Xple* No synpalpi appeared in the F 1 or F 2 progeny* If the mutation is on Chromosome II it would have appeared with Diohaete, since both homologues of Chromosome II were obtained from the mutant; or in the last individual, ? Xple, whioh had both homologues of Chromosomes II and III derived from the mutant. If the mutation is on Chromosome III it would -.69 have appeared with Curly since hoth homologues of Chromosome III were ob-tained from the mutant, or in the last individual, ? Xple, as mentioned before. The only significant result was that the ratio of males to females in the ? Xple individuals was; 2 females: 1 male, { i i i ) Chromosome II In the test for linkage with Chromosome I I , Lobe Curly stock was used: Lj.Cy - each on a separate homologue of the 2nd Chromosome. On February 25 the following crosses were set up: A 1 Lobe Curly virgin? x 1 Xple orange synpalpi<r> B 1 Lobe Curly virgin ? x 1 Xple orange C 1 Lobe Curly virgin ? x 1 Xple synpalpi o-"> D 1 Lobe Curly virgin? x 1 Xple & E 1 Lobe Curly virgin? x 1 wild 0" 7 - control. The F results are l i s t e d in. Table XLIII. Table XLIII i B C D E (Control) Lobe oV 27 34 34 38 45 ?? 35 41 37 28 45 Curly o-v 36 32 40 25 41 ?? 38 36 32 24 42 The F 1 counts were as expected - 1 Lobe ? :1 Curly? :1 Lobe©'"': 1 Curly cr* . F 1 interses were set up consisting of a cross of 1 F 1 virgin Curly ? x 1 F 1 curly . The F 2 results are li s t e d i n Table XLI7. 4 -70 -Table XLIV Curly n Synpalpi cr'o- 7 Synpalpi A 154 143 1 cr1 92 '76 1? B 137 123 2 crV 7 69 56 0 C 150 128 4 o-V7 58 73 1 cr? D 165 161 1 cr? 33 34 0 E 139 151 0 72 71 0 (Control) ? U refers to the individual whioh has obtained homologues of the 2nd ohromosome from the mutant f l y . The expected -ratios in the F 2 counts were: 2 Curly:1 TE I f the mutant i s associated with the 2nd ohromosome i t should have appeared in the ?U individuals since they had obtained both homologues of the 2nd ohromosome from the mutant f l y . It i s not l i k e l y that the mutant, synpalpi, i s associated with the 2nd chromosome since synpalpi :occurred in Curly and US individuals - in fact most appeared in the Curly f l i e s . (iv) Chromosome III In the test for linkage with Chromosome I I I , Dichaete Gluecb stock was used: D/Gl - each on a homologue of the 3rd chromosome. On March 5 the following crosses were set up: A - 1 Dichaete Glued virgin ? x 1 Xple orange synpalpi or? B - 1 Dichaete Glued virgin? x 1 Xple orange cr7 C - 1 Dichaete Glued virgin? x 1 Xple synpalpio-7 D - 1 Dichaete Glued virgin ? x 1 Xple ^ E - 1 Dichaete Glued virg i n ? x 1 w i l d o 7 7 - control. The F 1 results are listed in Table XLV. - 71 -Table XLV A B G D E (Control) Diohaete 23 22 17 31 22 ? ? 25 26 19 29 17 Glued o V 27 22 17 33 19 ? $ 29 23 15 28 14 The P 1 counts were as expected: 1 Dichaete? :1 Glued? : 1 Diohaeteo-7:1 Glued F 1 crosses were set up consisting of 1 P 1 virgin Dichaete? x 1 P 1 Dichaete o-7 2 F results are listed in Table XLVI. Table XLVI Dichaete cr'cfl ?? Synpalpi Synpalpi A 108 176 0 58 89 1 B 120 127 0 63 62 1 cr? C 125 189 0 46 89 0 D 128 196 1 cr? 49 76 00 E 176 191 0 85 87 0 (Control) 7E refers to the individual which has obtained hoth • homologu.es of the 3rd chromosome from the mutant f l y . The expected ratios in the F 2 counts were: 2 Dichaete:1 ?N I f the mutant i s associated with the 3rd chromosome i t should have appeared in the ?N individuals since they had obtained both homo-logues of the 3rd chromosome from the mutant f l y . The F 2 results were very significant in that the ratio of males to females in both Diohaete and ?U individuals was approximately - 72 -1 male:2 females - (normal ratio i s 1 : 1 ) . This may he explained by the fact that one half the males received a Xple synpalpi or Xple X homo-logue from the original Xple synpalpi or Xple male - thus being homozygous for this mutant character. A combination of this 1 s t chromo-some with the 3rd chromosome which was heterozygous or homozygous for the homologues received from the original Xple synpalpi or Xple male parent proved lethal causing one half the males to die. - 73 -(v) Summary Prom the preceding tests for linkage with Chromosomes I, II and III, it seems very evident that the mutation is associated with Chromosomes I and III* In the test for Chromosome I synpalpi appeared in individuals whioh were homozygous for the Xple synpalpi X chromosome* In the test for Chromosome III a combination of this homozygous con-dition of Xple synpalpi on Chromosome I and Chromosome III, which is heterozygous or homozygous for the homologues received from the original Xple synpalpi or Xple male parent, proved lethal - causing one half the males to die* - 74 IV Discussion of Results The preliminary investigation and study of stocks brought forth many significant facts as to the nature of the mutant synpalpi. The mutant is not a gene mutation but more likely a chromosomal aberration, since Mendelian ratios were not observed to occur, and since the mutant appeared irregularly but persistently. A lethal factor is suggested by the small number of flies resulting from synpalpi crosses; the ab-sence or small number of synpalpi in the F 1 of~synpalpi crosses but its reappearance in 2-8$ of the P2 progeny; and by the death of flies in two different stages of development. It is assumed that the homozygotes die and the P 1 intersesof heterozygotes produce more synpalpi in the P 2 (b«y segregation and reoombination). The association of the lethal factor with the X chromosome is assumed beoanse few male synpalpi emerge. A deficiency of a part of the X chromosome would explain the death of the males and homozygous females. The possibility of such a deficiency was further substantiated by a review of the literature on deficient X chromosomes. One of the 55 deficiencies of the X ohromosome listed by Bridges and Brehme (1944), vz. Df(l)svr (Sutton, 1943), was found in 9 females from a cold treated mother. The Xple stock used in the present investigation and from which the synpalpi flies' arose was chilled in its transport to the laboratory from the supply house; there-fore, it is quite possible that such a deficiency of the X chromosome as that recorded by Sutton (1943) could have occurred. The synpalpi flies have most Xple characteristics - the marker locations, echinus (5.5), out (20.0), vermilion (33,0), garnet (44.4), 75 -and forked (56.7) being v i s i b l e ; therefore a very small deficienoy of the t i p of the X chromosome must be involved. Yellow (0.0), aoheate (0.0+), and soute (0,0 +•+ ) appear on the X chromosome in the order named - yellow being at the very t i p . The literature on a l l deficiencies in the region of yellow, aoheate)and scute was reviewed and many sim-i l a r i t i e s in results to those recorded in this investigation were noted. Df(l) 260-5. This deficiency, recorded by Demerec and Hoover (1936), involves a loss of the four terminal bands. There i s no pheno-typic effect and the females have a reduced f e r t i l i t y . Synpalpi f l i e s have a reduced productivity as seen in the various preliminary crosses. Df(l) 260-1. Sight terminal bands are deficient proving lethal in the egg stage (Demerec and Hoover (1936)). The injurious effect of this i s due to the absence of the locus 1J^, which i s of v i t a l importance to the organism. This d.ctWal may account for the low F 1 counts and the absence of dead larvae in the Xple crosses of the present investigation. Q Df(l) sc * Paulson (1940),. The deficienoy whioh i3 in the region of yellow, achaete, and soute affects the later embryonio stages. The tracheal system f a i l s to f i l l with a i r - corresponding to the defective tracheal system found in pupal dissection i n this synpalpi investigation. The deficienoy i s also recorded by Sturtevant and Beadle (1936) as affecting the males deficient for yellow and aoheate causing death in the egg stage. Df(l) 260-2. The literature on this deficiency proved the most valuable and applicable to the mutant under investigation. As recorded - 76 by Demerec and* Hoover, (1936) eight terminal bands are involved, which, when homozygous deficientjare lethal. The females with the deficiency ' die in the late pupal stage and the males in the egg 'stage or i n larval development. These ages of death correspond to the two ages found 'in the dissection of pupae oases of the mutant synpalpi stock. The larval period i s considerably lengthened in Df(l) 260-2 as i t i s in the case of synpalpi crosses. Df(l) 260-2, recorded by Sutton (1943), i s lethal in both homozygous and hemizygous conditions - a fact which would account for fewer male synpalpi appearing. The deficiency affects yellow and achaete but not soute and there i s a considerable variability in the occurrence of the deficiency - both facts also applicable to the synpalpi stooks. Kaliss, Df(l) 260-2, (1939) reported that no males deficient for this region are viable. The total mortality of this stock i s 33.4% -death occurring at egg, larval, and pupal stages: 20.6 - 31.1$ as eggs. This high percentage of death in the egg stage would account not only for the low counts from synpalpi crosses but also for the absence of dead larvae in the examined cultures. The possibility that from 3 - Bfo of the deficient eggs hatch may indicate that the synpalpi, which appear in this percentage range, are those deficient f l i e s that do emerge. Kaliss also stated that the deficient larvae have tracheae which f a i l to f i l l with air and also a feebleness of motion associated with a lower metabolic rate. This may explain the i n a b i l i t y of the synpalpi f l i e s to emerge. Heat and cold treatments proved that the Xple stock i s very sensitive to variations from the optimum temperature of 25° C. - 77 -Higher (29° C. and-35° C.) and lower (15° C.) temperatures markedly de-creased the v i a b i l i t y so that few f l i e s emerged from the treated stocks. Those that did emerge were normal and produced only normal offspring, therefore, i t i s assumed that the experimental temperatures increased the lethal effect of the deficiency. This i s borne out by the fact that the treated cultures contained numerous dead pupae in two distinct stages of development as described previously. : In cold treatments an exposure at 4-5 and 5-6 days after development began proved most c r i t i c a l - no f l i e s emerging from these cultures. Demerec and Hoover (1939) i n their investigation of Df(l) 260-2 reported that i n the larvae a l l deaths occurred before 5.5 days. In the test for linkage with Chromosome I, using yellow stock, synpalpi appeared .only in f l i e s which were homozygous for the Xple synpalpi chromosome received from the synpalpi male parent. In the C1B -test oross synpalpi occurred only in the males which were hemizygous for the X chromosome but not in the females whioh were heterozygous. This again indicates the significance of Chromosome I in the inheritance pattern of the mutant. The absence of lethality in the C1B test crosses, where brother sister matings were made, might be explained by the translocation of the deficiency onto one of the autosomes. The deviations from the expected ratios obtained i n marker tests with the third chromosome, suggests that Chromosome III i s probably associated in some way with the produc-tion of the synpalpi mutation. The mutation i s not a simple deficiency of the X chromosome. - 78 -7 Summary 1. The inheritance pattern suggests a chromosomal aberration rather than a gene mutation* 2. The evidence would seem to point to a deficiency involving a few terminal bands of the X chromosome* 3 . The mutation i s not a simple deficiency of the X chromosome alone, but appears to involve the third Chromosome as well. r- 79 -Literature Cited Braun, Weiner 1940, Experimental evidence on the production of the mutant waristapedial by a change of developmental velocities. Genetics, 25*143 - 149. (Biological Abstracts 1940:9867). Bridges, C. B. and K. S. Brehme 1944. The mutants of Drosophila melanogaster. Carnegie Institute of Washington Publication, 552, Washington, D. C. Demereo, M. and Margaret E. Hoover* 1936. Three related X-chromosome deficiencies in Drosophila. The Journal of Heredity, 27:206 - 212. Gordon, c. and J. H. Sang* 1941. The effect of environment on the exhibition of the mutant "antennalesa'* in Drosophila melanogaster. The Proceedings of the 7th International Genetic Congress, 131. Kaliss, E. 1939* The effect on development of a lethal deficiency in Drosophila melanogaster: with a description of the normal embryo at the time of hatching* Genetics, 24:244 - 270* Poulson, D. P. 1940* The effects of certain X-ohromosome deficiencies on the embryonic development of Drosophila melanogaster. The Journal of Experimental Zoology, 83:271 - 325. - 80 -Robertson, Charles William, 1937, The metamorphosis of Drosophila melanogaster - including an accurately timed account of the principal morphological changes. The Journal of Morphology, 59:351 - 399. Startevant, A. H. and 0. W. Beadle* 1936* The relations of inversions in the X chromosome of Drosophila melanogaster to crossing over and disjunction. Genetics, 21:544 - 604* Sutton, E. 1943* A cytogenetic study of the yellow-scute region of the X chromosome in Drosophila melanogaster. Genetics, 28:210-217* Villee, Claude A. 1942. The effects of oold treatments on the development of the mutant Maristapedial Genetics, 27:173. 1944. The effects of temperature on the expression of "aristapedia". The Journal of Experimental Zoology, 93:75 - 98. (Biological Abstracts, 1944, 19499). Plates All the figures are photomicrographs of the heads of living Drosophila melanogaster at a magnification of 60 - light source as indicated* - 81 -Plate I Head of a normal Xple fly, dorsal view, showing the normal position of the antennae. Light source dorsal* Head of a synpalpi Xple fly, dorsal view, showing the protrusion of the forehead1 and the fusion of the antennae. Light source dorsal* 82 -Plate I Fig. 2 Plate II Fig* 3 Head of a normal Xple fly, dorsal view - similar to Pig, 1* Light source ventral* Pig. 4 Head of a synpalpi Xple fly, dorsal view - similar to Fig. 2. Light source ventral. Plate I I -85-Plate III Head of a normal Xple fly, ventral view, showing the normal position of the antennae. Light source dorsal* Head of a synpalpi Xple fly, ventral view, showing the fusion of the antennae. Light source dorsal* - 86 -Plate IV Fig. 7 Head of a normal Xple fly, ventral view, similar to Fig. 5 . Light source ventral. Fig. 8 Head of a synpalpi Xple fly, ventral view, similar to Fig. 6 . Light source ventral. - 8 8 Fig. 8 - 89 -Plate 7 Fig. 9 Head of a normal Xple f l y , lateral view, showing the position of normal antennae. Light source dorsal* Fig* 10 Head of a synpalpi Xple f l y , lateral view, showing the protruding characteristic of fused antennae* Light source dorsal* 90 -Plate 71 Pig. 11 Head of a normal Xple fly, lateral view, similar to Fig. 9* Light source ventral. Fig. 12 Head of a synpalpi Xple fly, lateral view, similar to Fig* 10* Light source ventral* - 92 

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