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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.  KAJffWar  BY MARGARET COLE  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF ARTS IN THE DEPARTMENT OF BIOIOGY AND BOTANY  THE UNIVERSITY OF BRITISH COLUMBIA APRIL, 1947.  ACZHOTSLEDGaiEin? S  The writer wishes to express her sincere appreciation to Mrs. Ruth Fields Brink. Assistant Professor, for a l l the assistance given during the following experimental study. Mrs. Brink* who discovered the mutation under investigation, was always ready to advise and showed keen interest and kindness at a l 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 A  II  1  Origin of Synpalpi  1  B Statement of Problem  1  Description of Mutant Stock under Investigation  III Experimental Study  2 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  (ii)  Discussion of results  3* Temperature treatments  19 24  (i)  Cold treatments  25  (ii)  Heat treatments  45  ( i l l ) Discussion 4»  6  Investigation of the l e t h a l i t y of the mutant  57 58  (i)  Grosses of Zple and Xple synpalpi  58  (ii)  C1B method  61  ( i l l ) Discussion  64  5» Determination of the linkage group  65  (i)  Chromosome I  65  (ii)  Chromosomes I I and I I I  68  Page ( i i i ) Chromosome I I  69  (iv) Chromosome I I I  70  (v)  73  Summary  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 i n the Xple Stock obtained from the B i o l o g i c a l Supply House, Chicago, i s so named because of the fusion of th£ antennae along the midline, a l l other Xple c h a r a c t e r i s t i c s being normal except f o r a s l i g h t protrusion of the forehead.  In stock cultures the mutant appears i n  greater numbers from normal Xple crosses than from synpalpi Xple crosses - with a predominance of female synpalpi.  Experiments were  carried out i n order to determine the nature of the mutation, the type of inheritance involved, and the linkage group with which the mutation i s associated. The mutant i s not a gene mutation but a chromosomal aberration since Mendelian r a t i o s were not observed to occur and since the mutation appeared i r r e g u l a r l y but p e r s i s t a n t l y .  A l e t h a l factor  is suggested by the small number of f l i e s r e s u l t i n g from synpalpi crosses; the absence or small number of synpalpi i n the F p a l p i crosses but i t s reappearance i n 2 - Qf> of the F  of syn-  progeny;  and  by the death of f l i e s i n two different stages of development early and late pupal stages.  The association of the l e t h a l factor  with the "X" chromosome i s 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 p o s s i b i l i t y of a  deficiency of a few bands at the t i p of the X T,  n  chromosome was  sub-  stantiated by a review of the l i t e r a t u r e on deficient "X" chromosomes. It i s believed that synpalpi f l i e s are those d e f i c i e n t 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  t o the head i n i t s e f f o r t s t o  e x t r i c a t e i t s e l f from the pupa case ( i t s tracheae b e i n g a t t a c h e d p o s t e r i o r l y t o the c a s e ) . The X p l e s t o c k i s v e r y s e n s i t i v e t o v a r i a t i o n s from o  optimum temperature  o  of 25 C.  the  ©  Temperatures of 35 C., 29  C,  and  15 C. i n c r e a s e d 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 s y n p a l p i a p p e a r i n g i n the F  or F  of t e m p e r a t u r e - t r e a t e d 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 t o l-fce a s s o c -  i a t e d w i t h chromosomes, I and I I I - t h e r e f o r e the mutant i s not a s i m p l e d e f i c i e n c y of the "X" the t h i r d chromosome as w e l l .  chromosome a l o n e but appears to i n v o l v e  The Study of a Possible New Mutation, Synpalpi, Occurring i n Drosophila Melanogaster  I Introduction  A. O r i g i n of Synpalpi On January 28, 1944,  a culture of the multiple stock, Xple,  was  received, i n a v i a l , by Mrs. Bath F i e l d s Brink from the General Bio l o g i o a l Supply House, Chicago. were placed i n a culture b o t t l e . acters.  The v i a l contained several larvae which Six f l i e s emerged showing Xple char-  These f l i e s were mated interse and on February 23 a mutation  appeared i n 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 t h i s 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 i n 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 v e r t i c a l l y 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 p u l l i n g of the antennae when they fuse. Comparisons of normal and synpalpi f l i e s , dorsal and ventral views to i l l u s t r a t e these points, are  shown i n 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 o r i g i n a l protruding position. This mutation i s very easily recognized as seen i n the plates named and i n a l a t e r a l 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  i s abnormally pointed and appears squeezed. The significance of this w i l l be mentioned i n a later discussion.  Ill  Experimental  Study  A.  Methods of Culture  The c h i e f culture requirements of Drosophila, oommonly c a l l e d the f r u i t f l y , were supplied as follows$  adequate food conditions,  l i v i n g yeast c e l l s whioh form the c h i e f food of the larvae, an optimum temperature of about 25° C ,  and s u f f i c i e n t  moisture.  The adequate food conditions were met by a culture medium whioh provided a good p h y s i c a l substrate and a s u f f i c i e n t 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 b o t t l e s , previously corked with cotton and s t e r i l i s e d , u n t i l they were approximately Vfliile the medium was  one-quarter f u l l .  s t i l l hot, small pieces of paper towelling were  placed i n the b o t t l e s to provide a place for deposition of eggs and a r e s t i n g place f o r larvae to pupate.  I f the cultures were to be kept  for a time Moldex-A was added to eliminate mold.  This was used i n  concentrates of 1 gram i n 100 c.c. of 95 per cent a l c o h o l .  Five c.c.'s  of t h i s solution were added to the medium five minutes before b o i l i n g was completed. 148 Lafayette St., New  Moldex -A was York C i t y .  final  obtained from Glyco-Products  Co.,  - 4The second condition, l i v i n g yeast c e l l s , was provided "by mixing a cake of Fleischmann* s Yeast i n a dropping 'bottle o f water and adding two drops of t h i s suspension to the food surface i n each culture bottle.  The yeast was not sown u n t i l the b o t t l e s were to be used.  Culture b o t t l e s were stored i n the r e f r i g e r a t o r therefore i t was necessary to allow them t o reach room temperature before yeasting. The optimum temperature of about 25° C. was maintained by placing cultures i n a thermostatically c o n t r o l l e d incubator. The oultures were kept s u f f i c i e n t l y moist by p l a c i n g a pan of water i n the incubator. F l i e s to be mated were added to the yeasted culture b o t t l e s . Since these f l i e s were under ether, i n order to insure that they would not f a l l on the medium and so damage t h e i r wings or not be able to detaoh themselves, they were brushed into a paper scoop and caref u l l y placed on the paper i n the b o t t l e . Etherization of the f l i e s was c a r r i e d out by.shaking them from t h e i r culture bottle into a b o t t l e , the a i r of whioh was saturated with ether.  Such a bottle was c a l l e d an e t h e r i z i n g bottle and was f i t t e d  with a cork which had a piece of cotton attached.  To saturate the a i r  of the b o t t l e , ether was poured on the cotton and the cork f i t t e d t i g h t l y into the b o t t l e .  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 f o r establishing cultures. V i r g i n females were secured from stock cultures by f i r s t shaking off a l l f l i e s . virgin.  The females which emerged 12 r 15 hours l a t e r were  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  =  c d - cardinal - formerly called orange. Appeared 2  i n chromosome carrying non-lethal Ins. (2L 4-2R)P. Eye color yellowish vermilion - O c e l l i 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 f u l l y later. The stock used to test for linkage group, Chromosome I , was yellow: y:  yellow - Wallace, 1 - 0.0 Body color r i c h yellow - hairs and b r i s t l e s 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]_.  - 6The stock used to test for linkage group, Chromosomes II and III, was Star dichaete: S/Cy D /ln(3L)PMe. 3  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^ D : Dichaete - Plunkett, 3* Inseparable from In(3L)P. 3  3  Wings extended and lifted at 45 degrees from the 6  body axis and elevated 30 degrees above* Alula gone* Viability of D /+ is better than that of T)/4-. D /D 3  3  and D /p are lethal* EEgA. 3  3  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 l i t t l e 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 l e s s viable. Best used as recessive,  RK^.  Cy: Curly - described previously. The stock used to test for linkage group, Chromosome III, was Dichaete glued: 3 D : Diohaete  3 ' -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 i n stock culture. The mutant, Synpalpi, appears persistently in the Xple and Xple orange stock cultures. In the Xple stock i t 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 s t 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  ??  Nov. 4 15 22 5 12 12 6 2 4 1 7 Total 30 38  Synpalp 1  1 (died)  1  ?? 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 count 1  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 20 .  In Bottle  1  #23 a cross was set up of the F synpalpi ? z F* normal o \ 1  F results 2  are listed in Table II. The abbreviation B for Bottle will be used hereafter. Table II  B #21 Nov. 15 20 22 Total B #20 Nov. 22 25 26 27 Total  Nornsal Sym >alpi ?9 ?? 2 6 3 4 4 8 9 16  12 5 14 18 49  13 10 10 19 52  2 1 0 3  2 4 1 6 1  Non nal Synpalpi B #22 Nov. 15 18 20 22 23 Total Totals of a l l Fl interses  99  B #23 Nov.- 18 20 22 Total  1  1 5 4  2 7 3 3 16  0  2 1 2 5  68 86  6  9  10  Nornial aV 3 2 1 6  3 4 2 9  •  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 and F . TABLE III BOttle Set Up Small Eggs Pupae First F Larvae or F NO. 2 Oot. 24 Oot. 26. Oot. 27 Oot. 30 Nov. 2 20 Oot, 30 NOV. 1 Nov. 2 Nov. 5 Nov. 8 21 Nov. 4 Nov. 5 Nov. 6 Nov. 10 Nov. 13 22 Nov. 5 Nov. 6 Nov. 8 Nov. 12 Nov. 15 20l Nov. 9 Nov. 10 Nov. 12 Nov. 16 Nov. 18 23 Nov. 5 Nov. 6 Nov. 9 Nov. 13 Nov. 17 1  2  1  2  Total Length in Days 9 9 9 10 9 12  -10In a cross of synpalpi o  x synpalpi ?  3  one would expect to  get a high percentage of synpalpi F - but the results showed only 2  approximately 9% synpalpi in F .  It is noticeable in the results in  2  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 accounted  for by a lethality of this synpalpi mutant - since the male  has only one X chromosome. The F results from the cross of P 2  $ x  1  synpalpi  also seemi to indioate a lethality character of the  normal  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: F  1  Cross of 2 Xple orange synpalpi ?? x 3 Xple orange oV.  results are listed in Table 17 - along with F results of B #30 1  parents from B #3. TABLE IV B #3  Nov. 4 5 6 7 Total  Nor ooal Synj>alpi B #30 CrV ?? ?¥ Nov. 9 29 23 9 7 1 12 1 4 13 3 1 15 42 35 0 1 Total Total F Count  Non ooal oV ?? 3 4 10 7 6 1 4 4 23 16  F  2 0  2  0  3  1  51  65  F* interses were set up - B #31, 32, 30 , 30 . synpalpi crosses* B #33 1 P synpalpi $ x 2 F^ c?if B #30 » 2 F synpalpi 5 £ c/V 1  1  Synj>alpi oV ?*  9  2  1  1  3  - 11 F  results are l i s t e d i n Table 7.  2  Table 7 Honnal  ??  B #31 HOT.13 1 1 1 15 3 3 18 3 1 20 4 4 22 2 3 Total 13 12 1 B fSO Nov*18 6 3 20 5 2, 3 22 1 10. 25 30 18 1 27 21 16 Total 63 49 4 1  ??  Nonsal  Nor coal Synpa l p i ¥9 2?  Synpa l p i  B #32 Nov*18 12 11 20 6 7 22 6 2 25 1 3  25 23  0  B #30 1 Nov.25 2 2 27 3 2 29 4 5 3 2 1 1 Seo. 2 10 11 5 Total  1 1  0 Total 3  2  B #30 Nov*25 4 27 8 5 29 Total 17 B #33 0 NOV*18 2 20 14 22 10 25 3 29 1 Total Total l 1 Interses 111 2  Synpii l p i  cry 99  5 17 5 27  P  0  0  4 11 13 5 33  1 2 3  0  95  7  6  Length of l i f e h i s t o r i e s are given i n Table 71 Table 71 Bottle No*  Set Up  Eggs  Small Larvae  Pupae  3 30 31 32, 30 30 33 30  Oot.22 Oot. 28 Nov. .4 Nov* 5 Nov* 9 Nov.15 Nov. 5 Nov.12  Oot.24 Oot.29 Nov. 5 Nov. 6 Nov.10 Nov. 16 Nov. 6 Nov.13  Oct.25 Nov. 2 Nov. 6 Nov. 8 Nov. 12 Nov.18 Nov. 8 Nov.15  Oot.28 Nov. 6 Nov. 9 Nov. 12 Nov.16 Nov.22 Nov.15 NOV.20  1  3  2  F  1  First_ or" F  8  Total length In Days* 9 11 9 8 9 10 13 13  NOV* 1 Nov. 8 Nov.13 NOV.13 Nov*18 Nov.25 Nov.18 Nov*25  In t h i s oross of Synpalpi? x normalcfithere was a more balanced number of males and females*  Synpalpi appearing as a result of F i n 1  terse constituted only 6% of the t o t a l F . The cross of F synpalpi $ 2  x.P  1  1  normal o^yielded only 3$ synpalpi - a l l males. There was also a  very poor r a t i o of males and females*  A comparison of Tables III and 71 shows that the life history of crosses of F synpalpi? x  normal a increased in duration - being  1  71  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:  F  1  Cross of 5 Xple orange $?x 2 Xple orange synpalpicr^  results are listed in Table, 711 • Table 711 B #5 Nov. 8 9 12 13 Total  Normal  oV  Synpalpi  ??  1 3 1  4 3 3 1 11  5  1 1  0  An •F^ interse was set up in B #51. F**" synpalpi cross: B #52 - F synpalpio^x F . normal? • 1  F  2  1  results are listed in Table T i l l . Table T i l l B #51 NOV. 20 22 25 27 Total  Nor mal  o^o^l  Synp*ilpi  ?$  30 27 5 12 11 8 46 47  '?* 1  0  1 1 1 4  B #52  Norrtttl  N0V.27 29 Deo. 2  cry  1*  11 2  3  1  Total  13  8  1  6  Synpialpl 9-9  Length of life histories are given in Table IX.  0  Table IX Bottle No.  Set Up  Small Larvae  Eggs  Pupae  Oot.30 Oct.31 -Nov. 2 Nov. 8 Nov. 9 Nov.13' Nov.13 Nov.14 Nov.17  5 51 52  The F  1  First F  Nov. 5 Nov.18 Nov.21  1  or F  Novi 8 Nov.20 Nov.27  2  Total Length In Days 9 12 14  count (Table VII) was very low considering that five  females were used. Males constitute only one-third of the t o t a l normally should be one-half. Both these facts suggest again the l e t h a l i t y of the mutant synpalpi. The F  2  count (Table VIII) - result of F  4$ synpalpi - a l l female.  interse - showed only  1  Cross #2 - reciprocal of t h i s - gave 6%  synpalpi - but both male and female appeared. As i n 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 w i l d $ ? x 2 Xple orange synpalpi Oo^. F  results are l i s t e d i n Table X.  1  All F  1  f l i e s were wild type.  Table X B #4 Nov.  Total An F F  2  1  9 12 13 15  Non nal dV ?? 5 9 5 13 1 4 2 3 25  17  Synpa l p i oV ??  0  interse was set up - B #41 and  results are l i s t e d i n Table XI.  0 #42.  - 14 Table XI Nor mal ?? B #41 Nov. 22 25 27 29 Total  13 10 1 1 1 4 1 4 16 19  B #42 Nov. 25 27 29 Total  18 10 8 36  19 13 12 44  Total of F^- interse  52 63  1  WiM Synpji l p i oV"> $?  Xple Orange Synpalpi Non aal ¥$ o V ?<? o V  0  1 1  0  0  0  0  1 2 2 5  0  0  0  0  0  6  0  0  0  0  0  Length of l i f e h i s t o r i e s are given i n Table X I I . Table XII Bottle NO.  Set Up  4 41 42  Oot.30 Nov. 9 Nov.12  Eggs Nov. 1 Nov.10 Nov.13  Small Larvae  Pupae  Nov. 3 Nov.13 Nov.15  Nov. 6 Nov.16 Nov.20  First F or F 2  Nov. 8 Nov.21 * Nov.23  1  Total Length In Days 9 12 11  No synpalpi appeared either i n F* or F . Cross #5 -Reciprocal of #4 B #6 - Cross of 1 Xple orange' synpalpi + x 4 w i l d ^ t y . results are l i s t e d i n Table X I I I . A l l F f l i e s were wild type. 2  7  F  1  1  Table XIII Non^ 9 B #6 4 Nov.15 6 6 18 0 6 10 Total  i Ipi 0  - 15 -  P P  2  1  interses were set up - B #61 and #62.  r e s u l t s are l i s t e d i n Table XIV. Table XIV  B #61 Nov.29 Dec. 2 4 Total  Wild Synpa l p i  Non nal ?? .  ?*  Xt>le Oranse Synp a l p i Nor mal <?$ orVl  3  4 4 3  4 1  11  8  8 5 1  0  0  0  0  0  B #62 Nov.27 Dec. 2 4 Total  14  9 6 2. 17  Total of F! interse  25  25  0  0  0  0  0  1  0  0  0  1  0  0  0  1  Length of l i f e h i s t o r i e s are given i n Table XV  Bottle No.  Set Up  Eggs  Small Larvae  Papa©  6 61 62  Nov. 5 Nov. 15 Nov.15  Nov. 6 Nov.16 Hov.16  Nov. 8 Nov.18 Nov.19  Nov.11 Nov.25 Nov.24  First F 2 or F Nov.14 Nov.29 Nov.27  1  Total Length In Days 9 14 12  Cross #4 and i t s r e c i p r o c a l , #5, had extremely low F* counts as seen i n Tables X and XIII - no synpalpi appearing.  Cross #4 P* f e -  males were i n the minority as compared to Cross #5 where the males were i n the minority. The F  2  counts o f both crosses - Tables XI and XIV -  were also very low, Cross #5 having the lowest.  No synpalpi appeared i n  - 16 P  2  progeny i n either  - t h i s may he explained by low counts and  G r o s s  l e t h a l i t y of mutant.  Cross #5 which had the lower F  longer l i f e h i s t o r i e s o f F  1  count also had  2  interse crosses - as seen i n Tables XII and  XV. Cross #6 B #300 - Cross o f 2 Xple synpalpi ? ? x 4 Xple s y n p a l p i c r ^ F  1  r e s u l t s are l i s t e d i n Table XVI. Table XVI B #300  F F  2  Nor coal or>cr  99  Nov. 9 12 13  3 10 8  1 13 6  Total  21  20  Synpsi l p i  99  0  0  interses were set up - B #301 and #302.  r e s u l t s are l i s t e d i n Table XVII. Table XVII  B #301 Nov.22 25 27 29 Total Total o f p l interse  Hor coal • cr?cr] 99 1 4 8 15 5 3 3 6 25 20 43  43  Synpa l p i  B #302  Hon nal  1  1  1  1 1 3  Kov.25 27 ' 29 Dec. 2 Total  6 6 2 4 18  4  4  92  9? 6 9 1 7  23  Synpa l p i 1 2 1 3  Length of l i f e histories are given i n Table XVIII.  1  - 17 -  Table XVIII Bottle Ho.  Set Up  Sggs  Small Larvae  Pupae  300 301 302  Oot.30 Nov. 9 Nov.13  Nov. 1 Nov.10 Nov. 14  Nov. 3 Nov.12 Nov.16  Nov. 6 Nov.18 Nov.20  First gF or W  1  T o t a l Length In Days  Nov. 8 Nov.21 Nov.23  9 12 10  Comparing FA r e s u l t s o f Cross #1 (Table I) Xple orange - the F  1  count i n t h i s cross (Table 2SVT) was lower and had no synpalpi as  Cross #1 had. The F Cross #1.  2  count showed 9fa synpalpi - the same as that o f  Length of l i f e h i s t o r i e s o f Cross #1 (Table III) and Cross #6  (Table XVIII) were also s i m i l a r . Cross #7 , B #100 - Cross of 2 Xple synpalpi F  1  5 Xple o ^ ,  r e s u l t s are l i s t e d i n Table XIX. Table XIX  F F  2  1  B #100  Nontnal  Oct. 25 28 Total  Synj>alpi  17 13  9?  o-"Vr> 9?  17 13  30  30  0  !  0  interses were set up - B #101 and #102.  r e s u l t s are l i s t e d i n Table XX. B #101 Nov. 4 5 6 7 9 12 13 Total  Norna l 2  9 2  2 1 1  5$P"  1  Nov. 6 7 8 12 13 Total  1  Total of F l interse  1  6 1 2  2 1 2  11  9  1  o  Table XX B #102  29  1 2 10 18 6 37  0  2 1 5  40  46  1  6  3 8 14 4  l i  Length of life histories are given in Tattle XXI. Table XXI Bottle  Eggs  Small Larvae  Pupae  Oct.11 Oct.13 Oct.25 Oct.26 Oct.28 Oct.29  Oct.16 0ct.28 Oct.31  Oct.19 HOV. 1 Nov. 3  Set Up  HO..  100 101 102  First F or F  1  2  Total Length In Days  Oct.22 Nov. 4 Nov. 6  11 10 9  Synpalpi occurring in F constituted 8% of total - higher than 2  that of the similar cross of Xple orange (Cross #2). In Cross #2 synpalpi were males - as contrasted to this cross, where synpalpi were mostly females. Cross #8 - Reciprocal of #7* B #200 - Cro ss of 1 Xple ? x 1 Xple synpalpi cr . 57  F results are listed in Table XXII. 1  Table XXII B #200 Nov. 8 9 12 13 15 Total F  1  Normal ?9 2 3 6 5 6 5 6 7 5 6 25 31  Synpalpi  cr>cr>  99  1 1 1' 1 3 1  interses were set up - B #201, #202.  Crosses were also set up of F synpalpi'. B #203 = 2F synpalpi cf^k P* 1  normal ?  . B #204 s F synpalpi o/* x F synpalpi ? - died. 1  F results are listed in Table XXIII. 2  1  1  - 19 -  Table XXIII B #201 HOT,22  25 27  Honnal c/M S? 21 18 13 9 2 2 36  29  T o t a l of all F interses 91  82  Total  Synpa l p i B #202 Nor ml Synp a l p i B #203 99 oVj ?? oVI %? 2 4 Nov,22 6 Nov.25 7 1 27 25 11 29 3 27 30 31 9 1 Deo* 2 29 12 4 Total 55 53 1 4 Total  Nor mal Syn] jalpi  oV| 4 11 9 7 31  f ??  9 3 15 4 31  1 1  1  1  8  Length o f l i f e h i s t o r i e s are given i n Table XXIV* Table XXIV Bottle NO.  Set Up  200 201 202 203  Oat.28 NOV. 8 Nov.13 Nov.13  Eggs  Small Larvae  Oct .30 Nov. 1 Nov. 9 Nov.13 Nov.14 Nov.16 Nov .14 Nov.16  Pupae  First F Of F 2  Nov. 4 Hov.18 Nov.18 Nov.19  Nov. 8 Nov.21 NOV.21 Nov.22  1  Total Length In Days 11 13 8 9  •  In the F  2  oount, synpalpi constituted 5$ o f the t o t a l .  The  Bimilar cross o f Xple orange (Gross #3) gave 4% synpalpi - which i s very near that o f t h i s cross (#8). Another s i m i l a r i t y of these two crosses i s the predominance of female synpalpi. N.B.  In a l l tables normal r e f e r s to normal antennae as opposed t o  fused antennae (synpalpi)*  (ii)  Discussion o f r e s u l t s .  The results o f the preceding crosses have been summarized and tabulated i n Table XXV. The time factor f o r the emergence o f F  1  f l i e s and F  1  counts  cannot be considered s i g n i f i c a n t 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 f e r t i l i t y  of the mutant synpalpi. Crosses #1 and #6, whioh were crosses of synpalpi 9 x synpalpi a produced 9* synpalpi in the F in equal numbers.  2  - synpalpi appearing in males and females  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 i s 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 than in crosses #1 and #6, 2  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  males and females*  2  In the case of Cross #7, Xple synpalpi? x X p l e O ' , where no synpalpi 7  were produced in the F , there was a predominance of female synpalpi 1  in the F . 2  In crosses #3 and #8, where the mutation was introduced by the male, there were fewer synpalpi in the F than there were in Crosses #2* 2  #7, #1 or #6* - the female synpalpi flies being i n greatest numbers* These crosses (3 and 8) also produced 6% synpalpi i n the F with a 1  greater predominance of male synpalpi* The fact that fewer synpalpi appeared in the T  1  than in the F  2  may indicate that most homozygotes die in the F and the heterozygotes, 1  by segregation and recombination, produce more synpalpi i n 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 a l l the similarities in ratios obtained*  Summary of the Preliminary Investigation* dross No,  Constitution  Average No. of P No, of NOo of No. of pJSyn- pl..Sjn- P Syn- Length p a l p i p a l p i p a l p i of l i f e history F 9? of F interse  No. of No. of % pl Syn- P l Syn- Synpalpi palpi  2  x  1  1  1  2 Xple orange synpalpi'- 79 x 3 Xple orange synpalpic  2.5  1  1  99  10 days Interse: I69 74 95 9 p l Syn<? ) r x: P l <r> ) 6 cf> s 9 Interse: 219 9 " 118cr>l01? l t 0  2  2 Xple orange synpalpi^ 119 2.5 x 3 Xple orange cr> 65o">54?  3:  0  p l Syn? )  1  Q  q  9  9  6  0  0  0  6 ,-•  6  7.  3  0  3  4  0  0  1  0  0  0  0  49 cr 60$ Interse: 97 4 46o-->51? F ?* ) 5 P Syn a*) 14. <r> 8 9 Interse: 121 0 (Wild: 520*63?) (Xple orange 6oj 1  35  5 Xple orange ? 17 x 2 Xple orange synpalpi* ^6o- ll9  6.0  0  1  12 '»  7  1  2  2  1  42 .2 wild 9 . x 2 Xple orange synpalpi ^(wild) 25o^.7? 1 Xple orange synpalpi R 16 5 (wild) x. 4 wilder* 6cr>10?  4  <  0  0  0  11.5  0  0  0  13  11  "  Interse: 51" 0 (nmd:25<^259; (Xple orange lcr?  TABLE XXV (Continued) Cross No.  Constitution  NO Of Average No. of Length F SynP SynSynof l i f e palpi palpi • palpi pi h istory cr>cf> 9? of F l Interse  No. of pi  6  2 Xple synpalpi^ 41 x 4 Xple synpalpio => 210^20$  7  2 Xple synpalpi? x 5 Xple synpalpio  8  60 1 Xple9 x 1 Xple synpalpio ^ 280^32?  60  -i  0  ,  0  6o0  e  x  ,  No. of P  2  1  No. Syn- of pi Synpalpi 99  No. of pi SynpaTji o-ly  0  0  11 days  Interse:  94  9  4  4  0  0  9.5 "*  Interse:  93  8  6  1  Interse: 182 92 cr> 90 9 pi 9 ) x P cr) ) 63 Syn )  5  8  1  2  1  0  1  35  11  "L  47 ori 47 9 41  o-  -7  52 ?  1  31 cf  1  32  9  o  3, Temperature treatments* Following the preliminary investigation of the type of inheritance an attempt was made to increase the penetrance of the mutation by prolonged cold and heat treatments* Villee (1942 and 1944) described 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 segmentation of the antennae disk is begun in the two day larvae along with the leg diso*  In normal flies i t 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 a l 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 a l l cold treatments. On December 18 at 1:15 p.m. the following orosses were set up ^* a l 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 cr  1  #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 againto 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 -  Nonmal Synpeilpi  #1  ?$  cry  Jan.20 1 1? 27 1 1? 0 Total  F  1  0  I  :  Normal Synp!alpi 99 °V| ?? Jan.27 32 26 29 8 7 Feb. 3 5 12. Total 45 45 0 0  Norraal SynpaIpi  #10  cry  99  "Jan. 27 35 32 Feb. 3 1 2 Total 36 34  .  #20  cry  9?  0  0  interses were set up as follows: #1 : B #1 #10t  : IF ? ? x IF  1  B #10 : SF^?? x 3F  1  1  1  #20: B #20  X  B #20  2  : 3F 9 9 1  - Jan. 27 - died Jan. 27  3F o-V - Jan. 27 1  x  : 3F 9 ? x 3F 1  7  Jan. 29  1  F oounts of the above F interses are listed in Table XXVII. 2  1  Table XXVII fio  Normsi l Synpsilpi #20 9? o-V <?? Feb. 10 Feb. 10 33 39 11 29 11 22 12 12 11 11 Total 66 79 0 0 Total 1  1  Non nal Synpsilpi cry  9 20 19 48  99  9?  15 31 12 58  l 0  l  #20  2  Feb. 10 11 12 Total  Normal Synptalpi ! o_ 99 4 9 6 15 11 10 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  -27-  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.  -  28  -  Dissection of papa oases: B #1  No larvae  v  Sampling of 22 pupa oases B#2  : Emergent: 34 oV'Z 99 Larval : 7  : Emergent: 21o&(i synpalpi) Larval : 9  No larvae 34 pupae  B #10  : Emergent: 110^0^(1 synpalpi) 4?? Larval : 19  No larvae Sampling of 19 pupa oases  B #20  synpalpi) 2 ? 9  : Emergent: 11 cr^S 94? Larval : 10  No larvae 31 pupae  B #70  ; Emergent: 19 Larval : 7  No larvae 24 pupae  B #60  : Emergent: 13 0 - ^ 3 synpalpi) 5 99 Larval : 1  No larvae 28 pupae  B #50  9£  No larvae 30 pupae  B #7  3  No larvae 44 pupae  B #6  : Emergent: 18 cryp3 synpalpi) Larval : 1  : Emergent: 11 c^Vl ? Larval ; 19  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 Jan* *6 7 8 10 Total  Normal Synpalpi #500 Normal Synpalpi #700 Nonoal Synpalpi Op 9? crV 99 99 OCT| 99 99 9? 38 37 4 4 Jan. 6 6 3 3 3 Jan. 6 11 18 5 12 7 12 0 0 7 1 0 1 2 7 3 1 2 5 8 0 2 0 0 7 4 0 0 8 0 0 0 1 . 10 14 16 0 1 9 10 0 2 61 63 4 6 Total 21 21 4 6 Total 14 19 7 18  #200 Jan. 6 42 40 7 11 15 8 7 7 10 3 4 Total 63 66  0 0 2 0 2  1 0 0 0 1  #600 Jan. 6 20 31 7 4 4 8 1 0 10 2 0 Total 27 35  2 7 0 2 0 1 0 0 2 10  B #300 - no eggs or larvae* orosses of controls were set up as follows: #100 : B #101 : F interse of normal-39 x S d - January 6, B #102 : F interse of synpalpi-2? x 2o->- January 8. 1  1  #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 interse of normal-3? x 3 o" - January 6, B #502 : F interse of synpalpi-29 x 2o*- January 7, 1  7  1  #600 : B #601 : F interse of normal-3?x 3 - January 6 B #602 : F interse of synpalpi-3 9 2er>- January 7 1  1  x  #700 : B #701 : F interse of normal-39* 3 or - January 6 B #702 : F interse of synpalpi- 3? x 3o-- January 8 B #703 : F synpalpi x F normal- 39 x 3o- January 8 1  1  1  1  F  2  7  1  results of controls are listed in Table XXIX.  7  - 30 -  Table XXIX For nal Synp.alp:  oV  %9  #101 Jan.l? 20 21 Total  14 16 23 21 17 19 54 56  0 0 0 0  #501 1 Jan. 17 2 2C 1 21 4 Total  #102 Jan*20 21 24 Total  13 17 14 14 26 28 53 59  2 0 3 5  #201 Jan.17 23 27 20 37 35 21 9 10 Total 69 72 #202 Jan,20 9 13 21 19 20 24 29 27 Total 57 60  Non nal Synpa l p i o-V> 9 ? o-Vi f¥  Normal Synp a l p i o-7f  orV  99  0 3 1 1 0 7 1 11  #701 Jan.17 21 20 17 21 11 Total 49  25 16 15 56  3 1 3 7  1 3 4 8  #502 2 Jan .20 12 15 1 21 17 18 2 24 21. 23 5 Total 50 56  0 2 1 3  2 1 2 5  #702 Jan.20 22 27 21 10 11 24 21 17 Total 53 55  2 0 4 6  3 1 5  9  0 0 0 0  #601 3 Jam. 17 21 23 2 20 13 14 1 21 21 20 6 Total 55 57  1 0 0 1  4 2 2 8  #703 Jan.20 21 24 Total  1 0 0 1  0 0 0 0  2 0 1 3  #602 3 Jan*20 18 1 21 16 0 24 21 4 Total 55  1 2 1 4  0 3 3 6  19 23 17 59  23 21 15 59  17 19 17 53  13 17 21 51  16 16 20 52  Additional oold treatments we're carried out i n order to include the age group 5 - 6 days i n the treatment. On January 24 at 12 a.m. the following crosses were set up: #1000 3 Xple synpalpi9?x 1 Xple synpalpi c r #2000 12 Xple  7  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.  -  31  -  The hot t i e s 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  B #2000  No larvae Sampling of 15 pupae  B #3000  : Emergent: 5 Larval : 3  7$  No larvae Sampling of 15 pupae  B #4000  69  : Emergent: . 9 Larval : 0  : Emergent: 4 c r 8 ? Larval : 3 7  No larvae Sampling of 15 pupae  : Emergent: 10 0 ^ 3 $ Larval : 2  The parents removed from bottles #1000, #2000, #3000, #4000 were placed i n bottles #1 , 2 , 3 , 4 , respectively on January 27 at 1  1  1  1  5 p*m* Another transfer was made on January 28 at 5 p.m. to bottles used i n heat treatment described l a t e r . The bottles were placed i n 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 results are l i s t e d i n 1  Table XXX.  -  32 -  •Table XXX n  Feb.17  1  Feb. 1 7  3  9  18  9  4  18  19  1  20  0  0  13  13  Total IP  &  Normal Synpalpi ?? °v ??  1  0  1  #4  Feb.17  47  19  53  27  20  15  15  85  91  1  8  19  19  4  20  20 2  F  0  0  Total  0  l(cued)  1  18  Total  2 normal  0  1  Feb. 1 7  1  18  1 ) to 1) normal  2 17  Total  0  Normal Synpalpi ? ? o V ??  13  (U  0  0  interses were set up:  1  .#lj : 3 F ?? x 3 F W February 1 7 1  #2*  : 3 F ?? x 3 F^Februsry. 1 ?  #2g  :  1  2F  1  (synpalpi-* normal)?? x  3  F V C A February  18  P results are tabulated in Table XXXI Table XXXI 2  IT ! 1  J.  Mar. 4 5 7  Total  Normal Synpalpi 9? o V ?? 5 4 16 25  #2i  Normal Synpalpi ?? 0 * 0 *  •ia  #2^  &  Normal Synpalpi crV ?v  6  Mar.3  10  16  Mar.3  3  4  9  3  4  8  3  19  5  11  10  5  3  4  7  8  6  7  1  0  Total  36  28  0  0  Total  38  35  0  0  24  16  23  0  0  The pupa cases containing dead flies were opened and examined. Observations. B $0- : No larvae 15  pupae  : Emergent:'  - part way out of oase) ( 4 -. palps not developed) 1 0 ? ? ( 7 - young-palps not developed. ( 3 - part way out of case) ScrVU  - 33 -  In these f l i e s i t was noted that the front of the head protruded as i n F i g . 2 - ventral view.  Pig. 2 - Ventral view of head of f l y i n pupa oase, showing protrusion of head. B #2  No larvae  1  A l l f l i e s emerged B l^  1  No larvae 2 pupae  B #4  1  : Emergent: 2?? palps folded down i(•palps* refers to antennae)  No larvae 94 pupae  : Emergent: 61otb^21 - forehead bulged palps protruding and w e l l developed. 35 - palps folded forehead not bulged - 3 of these show traoheae attached to posterior of pupa case. 1 - very young - head poorly formed - l a r v a l 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 together. 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 #1 , 2 , 3 , 1  1  1  4 were obtained by transferring the original parents to bottles 1  10000 , 20000 , 30000 , 40000 respectively - as described in the seotion 1  1  1  1  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 flOOOO  'Hotmal  1  Feb. 12 14 Total #30000 Feb. 12 14 Total  9?  40 48 11 13 51 61  fgoooo  Synj>alpi o V  99  0  1  13 10 0 0 13 10  99  0  23 37 36 32 59 69  0  0  0  #40000 Feb. 12 14 Total  65 67 21 24 86 91  0  0  0  interses of controls were set up: 1  - 3 P ?? x 3 F/W-  1  B #A  1  20000 ' •» 3F 99 1  X  3  I^cPcFL  - February 12  B #B - February 12  30000  - 3 F ?? x 3 f ^ o V - B #C  40000  - 3 . F ^ x 3 . ? o V - B f D - February 12  1  1  2  99  Synpalpi  1  B #10000  F  Normal  Feb. 12 14 Total  1  F  1  1  l  l  - February 12  ,  results of oontrols are listed in Table XXXIII.  - 36 .Table XXXIII A Feb.24 25 2? 28 Total  normal o-V ?9 0 0 4 5 31 30 23 17 58 52  C Feb.24 59 25 19 27 39 28 11 Total 128  52 24 26 11 113  Synpalpi cy<y 9%  i  1  0  1  0  0  1  0  1  B  normal  Feb.24 25 27 28 Total  2 9 4 10 12 11 11 32 30  D Feb*24 25 27 28 Total  16 14 54 4 88  99  22 12 60 5 99  Synpalpi 9? 0 1  0  l  1 0 0  l i 2  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 counts of wild oontrols: cry? ?9 77 fl 81 1  #10  85  80  #100  87  81  #1000  90  93  F* interses were set up and F total counts were: 2  #1  1  #10  1  #100  1  #1000  1  120  125  134  130  127  125  125  121  The results of the preceding cold treatments have been summarized and tabulated in Table XXXIV.  Table XXXIV - Cold Treatments Cross NO.  Constitution.  Age When Treated  1  13 Xple?? 4-5 days x 15 Xpleo*^  10  13 Xple?? 3-4 days x 15 Xple 6* a*  100  13 Xple?? Control x l 5 Xple 6*0" (25° C )  l No. of Synpi Emerged p a l p i 2 .0 F  Life History  Dec.18Jan.20 In cold Dec.23 Out Jan.13 0 70 Dec.19Jan.25 36^"34?? In cold Dec.23 Out Jan. 13 10 {8%) Dec.20134 Jan. 2  2 Synpalpi  D i s s e c t i o n of F-*- Pupae  P  No. of F  2  1  0  0  NO larvae Sampling of 22 pupae Emergent: 1800*3?$ Larval : 1  145 66°-V79 ??  0  No larvae 19 pupae Emergent: 13 (3 syn) Larval  Interse: 114  540V6O  ?£  :  1  4??(4#)  *  P syn. Interse: 122  10  580V64??  50*0*  1  2  14 Xple?? 4-5 days x l 5 XpleoV  Sinpa\p\  0  0  Dec.18 In cold Dec.23 Out Jan.13  0  (Q%)  5  0  ??  No larvae 44 pupae Emergent: 340^3?? Larval : 7  Cross No.  Constitution  Age When Treated  20  14 Xple ¥ ¥ x l 5 Xplecf^  3-4 days  14 Xple 2¥ x l 5 Xple^o*  Control (25° c )  200  Table XXXIV (Continued) pl No. of F* No* of Life SynHistory F palpi Emerged 90 45°-V45??  0  132 3 (2%) 65<yV67?? 2 o V l %  F^ Synpalpi  Dec.19Jan.25 In c o l d Dec.23 Out Jan.13  162 6 9 0 V 92??  1 V  Dec.20Jan. 2  Interse: 147 69«V72 ??•  692(3%)  P syn. Interse: 124  D i s s e c t i o n of P Pupae 1  No larvae 28 pupae Emergent: jqc?b»3$ Larval : 7 ,  0  1  2  12 Xple?? x28 Xple Co*  1  3-4 days  178  2??(#)  Jan.27Feb.17 In c o l d Jan.31 Out Feb.14  7  (#)  S O o V 6 4 99  30V4 $?  Interse: 73 3 8 ^ 35 n F syn.? x F 0^ 59  0  No larvae No pupae  1  1  0  360*0*25??  2000  20000  3  12 Xple** x 28 Xple o" a"  5 - 6 days  12 Xple ?? x28 Xplec^o  Control (25° C)  71  0  128  0  0  Jan.24 In c o l d Jan.30 Out Feb.12  0  Jan.31Feb.10  63  (  32 0 ^ 3 1 ? ?  0  19(2%)  No larvae Many pupae sample of 15 Emergent: 5oV7^ Larval : 3  Constitution.  Cross No©  1 Xple 4-5 oays Synpalpi?  3  3 Xple Synpalpi tt n  30 300 i  Age When Treated  3-4 days Control (2SO c)  3 Xple ?? 3-4 days Synpalpi  1  Table XXXIV (Continued) pi No. of F Life No. of SynHistory F Emerged p a l p i No eggs <>r larvae  ti  II  II  II  II  it  26  10000  1  5  50  lit  D i s s e c t i o n of F Pupae 1  II  13.0-V13??  0  5-6 days  0  0  Control (25° O  112 51cV6l¥¥  0  5 Xple 4-5 days orange ?-$ KlO Xple orange © V tt 3-4 days  2 Synpalpi P  it.  1 Xple Synpalpi 1000  2  Jan.27Feb.17 In cold Jan .31 Out Feb.14 Jan.24 In cold Jan.30 Out Feb.12  53  Jan.31Feb.10  111  Dec.19In cold Dec.23  0  0  0  No larvae 15 pupae Emergent: Larval :  5°V10?? 0  P  No larvae Many pupae sample of 15 Emergent: 9oV6?S Larval : 0  0  No larvae 24 pupae Emergent: l l o V 3 ? 2 Larval : 10  No eggs cr larvae  0  0  Cross No. 500  Constitution.  Age When Treated  Control 5 Xple orange 99 (25° C) x 10 Xple orange o-V  No. of pl  Emerged  Table XXXIV (Continued) l Life No. of P SynHistory palpi P  9 (18$ Dec.2051 25oV26?? 4 0 V 5 ?$• Jan. 2  1  6  5 Xple orange 99 x 11 Xple orange <rV  0  0  0  0  ti  3-4 days  600  tt  Control (25° C)  P  2  Synpalpi  D i s s e c t i o n of P Pupae 1  12 (9%) lcr>  119$  1  4-5 days  60  Interse 130 60oV709? P Syn. Interse 114 53crv61?? 0  2  Dec.18Jan.20 In cold Dec.23 Out Jan.13  Dec.-19 In cold Dec.23 Out Jan.13 12 (ie$) Dec.2074 290-V-45?? 2oV»10?? Jan. 2  0  Interse 121  8 (7%) 3<3*6» 5?$  0  No larvae 30 pupae Emergent: 21oMl syn) Larval : 9  0  No larvae 31 pupae Emergent: l l c V l ? Larval : 19  9 (7.30  560V65??  P Syn. Interse 118 59°-V59?? 1  10 {&fo ) 40^6 ?¥  Cross No.  4000  40000 7  Constitution  Age When Treated  8 Xple orange ?? x 10 Xple orange cr'cri  3-4 days  it  it.  2; Xple orange syn. ?9 x 4 Xple orange  5-6 days  Table XXXIV (Continued) pi No. of Life SynNo. of F History pi p a l p i Emerged 13  « o^i?^  1  4  0  Control (25° C)  177 860^91?$ 4-5 days. 0  0  0  0  It!  3-4 days  0  Synpalpi  D i s s e c t i o n of F-*- Pupae  0  0  No larvae 94 pupae Emergent: 61^0^30?$ Larval : 3  0  0  No larvae Many pupae - sample of 15 Emergent: l O o - i c 3?S Larval : 2 1  Jan.31— Feb.10  192 89o^l03??-  0  Dec.18In cold Dec.23 Out Jan.13  0  0  No larvae 34 pupae Emergent: l l c ^ l syn) 499 Larval : 19  0  Dec.19In cold Dec. 23 Out Jan.13  0  0  No larvae 41 pupae Emergent: 120*0(7 syn) 3 9S Larval : 26  syn. cr^cyi  ,70  Jan.27Feb.17 In cold Jan.31 Out Feb.14 " Jan.24In cold Jan.30 Out Feb.12  2  5 (2.90 4??  >oss No.  Constitution.  Age When Treated  700  2 Xple orange syn. ??• x 4 Xple orange syn. o-~V  Control (25° C)  3-4 days  3000  2 Xple orange syn. ¥9 x 3 Xple orange syn. cr^o ti  30000  tt  Control (25° c)  Table XXXIV (Continued) pl No. of F Life F* No* of SynSynHistory pi palpi palpi Emerged 24 ( 4 # ) Dec.20- Interse is m>) 120 57 7o->-a799 Jan. 2 7oV»8 9% 560V64??. 21o^36?? F syn. 15 (125?) Interse 123 6c^9 9 $ 59o-v6"4?$ 3p-">(ljO p l syn? X P"*- Cf) 52o^52 9 ? 2 o'V 0 0 0 Jan.27Peb.17 In cold Jan.31 Out Feb.14 Jan.240 0 0 0 In cold Jan.30 Out Feb.12 242 0 Jan.3123 19- (.50) 12 803114?? Feb.10 13o>b*L0?? 2  D i s s e c t i o n of F pupae  1  77  5-6 days  No larvae 2 pupae Emergent: 2 ? $ Larval : 0 No larvae Many pupae-sample of 15 Emergent: 4 O 0 V 8 9 $ Larval : 3  44  Cold Treatments Observations: In a l l 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 P  1  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 and F - in a l l oases the controls produced a fair percentage of 2  synpalpi (from 7$ to 12$).  1  - 45 (it) Heat treatments* Temperatures of 35° C* and 29° 0. were used in heat treatments* On January 26 at 5 p.m* the following crosses were set up: #10  - 3 Xple synpalpi ¥?x 1 Xple synpalpi o?  #20  - 12Xple¥? x 28 X p l e o ^ (from Xple synpalpi stook)  1  1  I30  - 2 Xple orange synpalpi??x 3 Xple orange synpalpio-V  1  #40* - 8 Xple orange?? x 10 Xple orange ^V?( from Xple orange synpalpi stock) On January 29 at 5 p.m. the parents were transferred to new bottlest  #10 to #100 * #20 to #200 , #30 to fSOO , #40 to I400 . 1  1  1  1  1  1  1  1  On January 30 at 5 p*m. the parents were transferred again to new bottles: #100 to #1000 , #200 to #2000 , #300 to #3000 , #400 1  1  1  1  1  1  1  to #4000* 1  On January 31 at 5 p*m. the parents were transferred to new bottles: flOOO to flOOOO , 1  1  §Z000  l  to #20000*, #3000* to #30000*,  #4000* to #40000* 1  On February 3 at 5 p.m* the parents were removed from bottles #10000*, #20000*, #30000*, and #40000**  These were used as controls.  On February 3 at 11:15 a.m. the bottles of series #10* 20* etc. (age, 5 - 6 days), #100 , 200 etc. (age, 4 - 5 days), and #1000 , 2000* 1  1  1  etc* (age, 3 - 4 days) were placed in the heat at 35° C. These bottles were removed from the heat on February 10 at 10 a.m. No F* flies emerged in any of the heat-treated cultures - a l l died in pupal or larval stages. The bottles were examined and the pupa oases were opened. Heisults of this procedure are listed below*  - 46 B #10  Ho larvae Sample of 12 pupae  B #20*  Emergent: Larval :  8o-V4?$ 0  Emergent: Larval :  8^3?? 1  3 larvae Sample of 12 pupae  1 B #30  No larvae Sample of 12 pupae  B #40  :  : Emergent: Larval :  0  No larvae Emergent: 2 cV"? Larval : 10  Sample of 12 pupae B JlOO  A few larvae (dead)  1  :  Emergent; 0 Larval : a l l  :  Emergent: 0 Larval : a l l  Pupae  :  Emergent: 0 Larval : a l l  B #1000*  A few larvae (dead)  no pupae  B #2000*  A few larvae (dead)  no pupae  B #3000  A few larvae (dead)  no pupae  B #4000*  Many larvae (dead)  no pupae  Pupae B #200  Some larvae (dead)  1  Pupae B #300*  No larvae or pupae visible.  B fiOO  No larvae  1  1  - 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 results, in Table XXXIII, 2  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 1 0 - 20* eto, numbers 10*, 1  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 72c Vand 76"?? - no synpalpi appearing. f  A l l heat treated bottles were examined. The observations are noted below: B #10*  No larvae 108 pupa cases : S mple of 15 a  Emergent: 6o-V74?$ Larval : 5 No larvae 120 pupa cases  : Sample of 15  Emergent :  Larval  : 6  ¥ Wb & f  <  - 48 #30*  No larvae 101 pupa oases : Sample of 15 Emergent : 6 trV4,9$ larval : 5 No larvae 115 pupa oases Emergent Larval 10 flies partly  #100^  : Sample of 15 : 9 0*6* 3 t 3 emerged  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  A  107 pupa oases : Sample of 15 Emergent : 6 o-V 2 9-S Larval : 7 No larvae 108 pupa cases : Emergent s Larval : #1000  A  77  No larvae 109 pupa cases t Emergent: Larval :  #2000^  Sample of 15 6 o^a 3 99 6  Sample of 15 4 0^0^ 1 ? 10  No larvae 119 pupa cases : Sample of 15 Emergent : 2 o-V" Z99Larval : 11 7  - 49 #3000'  No larvae  A  103 pupa oases Emergent Larval #4000^  Sample of 15 3 ofo-i 1 $  11  No larvae 126 pupa oases Emergent Larval  :  Sample of 15  i  3 crlcP 2 $  t 10  P results of controls are listed in Table XXXV along with the 2 2 1 F counts. F counts were the result of F interses. 1  Table XXXV #10000* pi  •g2  Nonsal o-">cr> ?? 63 67 50 51  Synpjilpi  pi 1  F  #30000^ F F  1  2  #20000| 2  Nonmal 99 75 71 37 48  crV  Synpalpi  99  3  #40000j 69 72 66 63  1  F F  1  2  63 82 67 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 c a r r i e d 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 i n heat at 3 5 ° 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 2 5 ° C. F and F counts are l i s t e d i n Table XXXVI - the F being the result of  #1 F  1  F  2  67  ' ?? 64  #3.0 72  ?$  counts  interses. #K 30  68 74  112 118, 121; 118; 98  69 1P7  Table XXXVI #1 000  «« 78  83  106 104  124  83  ££00  £0 70  78  127 117  ?¥  70  9?  80  #2C 100  73  89 123 105 111 130  92  85  135  A l l f l i e s were normal w i l d type - the heat treatments had no to effect other than^speed up l i f e h i s t o r i e s .  No f l i e s died i n l a r v a l or  pupal stages. A l l the r e s u l t s from heat treatments are summarized and tabul a t e d i n Table XXXVI.I. Observations: The Xple stocks were very sensitive to heat - temperatures of 29° and 3 5 ° C. k i l l e d a l l - so that no F  1  emerged.  synpalpi i n the controls was lower than u s u a l .  The percentage of  Cross No. IO*-  Constitution. 3 Xple syn.  Age When Treated 5 - 6 days  1 Xple syn. 100  1  1000  1  3  lococr  11  4 - 5 days  it  3 - 4 days  it  Control  Table XXXVII - Heat Treatments pi Temp. No., of No. of T Life p2 Syn- History C. F SynRmereed p a l p i palpi 0 Jan.280 0 0 35° In heat Feb. 3 Out Feb.10 0 0 0 0 Jan.2935* In heat Feb. 3 Out Feb.10 Jan.300 0 0 0 35° In heat Feb. 3 Out 2  25°  112  51<^61?  0  tt  5 - 6 days  290  0  0  i  ti  4 - 5 days  29°  0  0  1000} A  tt  3 - 4 days  29°  0  0  100  Jan.JL Feb.10 Mar«10 In heat Mar.17 Out Mar.24 Mar.12In heat Mar.17 Out Mari24 Mar.13-, In Mar. 17 Out Mar.24  111 590^2$  D i s s e c t i o n of F pupae No larvae Sample of 12 pupae Emergent: 8 c / V 4 ? $ Larval : 0 A few dead lapvae Sample of 12 pupae Emergent: 0 Larval : 12 A few dead larvae Pupae - 0  l°t'9#)  0  e  0  0  0  0  No larvae 108 pupae - sample of 15 Emergent: 6^4?$ Larval : 5 No larvae 105 pupae - sample of 15 Emergent: 4 o - V 4 9$ Larval : 7 No larvae 109 pupae -sample of 15 Emergent: AcrV'l ? Larval : 10  Table XXXVII - Heat Treatments (Continued) Constitution  3ross No©  Temp. No.-.of Age When P C. Treated Emerged  toooo^ 3 Xple syn.?? Control 25° 1 Xple syn.o'  130  ^ S y n - Life No« of p a l p i History P 9  0  Mar«,14«* 101 Mar.24 50o-^51??  630*67??  7  20  12 Xple  1  28 Xple  200  1  in  2000  1  11  20000  1  it  i  200*  D i s s e c t i o n of P pupae  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  4-5 days  35°  0  0  0  0  Some larvae Pupae: Emergent: 0 Larval : All  3-4 days 35°  0  0  0  0  A few dead larvae Pupae: 0  128  0  Jan.29In heat Feb. 3 Out Feb.10 Jan.30In heat - Feb . 3 Out Feb.10 Jan.31Feb.10  0*0*  Control 25°  59069$ 20  p2 Synpalpi 1^(1$)  it  5-6 day3 29°  0  0  it  4-5 days 29°  0  0  MaroilIn heat Mar.17 Out Mar.24 Mar.12In heat Mar.17 Out Man>^  63 1?(#) 32051? 0  0  0  0  No larvae 120 pupae - sample of 15 Emergent: ^cr'cy'^^ Larval : 6 No larvae 112 pupae-sample of 15 Emergent: 40^3?$ Larval : 8  Table XXXVII - Heat Treatments Cross No,  Constitution.  2000*  12 X p l e ? ?  Age Temp.' No.-.of When F C. Treated Emerged  2  Dissection of F pupae  0  No larvae 119 pupae - sample of 15 Emergent: 2<rV2 9? Larval : 11  29°  0  Control  25°  146  5-6 days  35°  0  0  Jan.28~ In heat Feb. 3 Out Feb.10  0  0  it  4-5 days  35°  0  0  Jan.29In heat Feb. 3 Out Feb.10  0  0  It;  3—4 days  35°  0  o . Jan.30In heat Feb. 3 Out Feb.10  28 Xple o-V  ti  20000*  2 Xple orange syn« ??  1  0) Mar.13In heat Mar.17 Out Mar.24 0 Mar.14Mar.24  No. of F SynF^ palpi  3-4 days  X  30  pi Life Syn- History palpi  (Continued)  X  3 Xple orange s y n o*cri  0  3 ? <A%) 75 37°'"48¥ 7  No larvae Sample of 12 pupae Erne rge n t : 9o-V 3 9 9 Larval : 0  0  300  1  3000  1  0  0  0  A few dead larvae  Table XXXVII - Heat Treatments  (Continued)  i  Cross No. 50000  x  Constitution. 2 Xple orange syn.  Temp. No. of Age When F C. Treated Emerged x  Control 2 5 °  23 13cU0?  Synpalpi  Life History  No. of p2  P Sympalpi 2  Dissection of F pupae  0  Jan.30Peb.10  0  Mar.11In heat Mar.17 Out Mar.24  0  Mar.12In heat Mar.17 Out Mar.24 Mar.13In heat Mar.17 Out Mar.24 Mar.14Mar.24  0  0  No larvae 107 pupae - sample of 15 Emergent: 6 o^cr>2 ?$ Larval : 7  0  0  No larvae 103 pupae - sample of 15 Eme r ge n t : 3 o ^ l ? Larval : 11  242 L28 o"' 114?? 0  7  1?(.S*)  X  3 Xple orange syn. o^o^ in  5 - 6 days 29°  0  it  4-5 days 29°  0  0  300C-1  It  3-4 days  0  0  50000*  t»  Control 2 5 °  141 690=72?  0  5 0 0  J  29°  0  No larvae 101 pupae - sample of 15 Emergent: 6 0 V 4 ? . Larval : 5 0  131  68o-">o-"> 63??  1S(.9*)  Table XXXVII - Heat Treatments (Continued) Cross No.  Age Temp. No...of P When C. SynF Treated Emerged p a l p i  Constitution.  1  X  8 Xple orange ??  5-6 days  35°  0  0  X  10 Xple orange cr' ' 0  7  Life No. of F SynHistory palpi F 2  Jan 28In heat Feb. 3 Out Feb.10  0  0  0  Dissection of F pupae No larvae Sample of 12 pupae Erne rge nt: 2 c r cr* Larval : 10 7  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 OOO  tr  3-4 days  35°  0  0  Jan.30** In heat Feb. 3 Out Feb.10  0  0  Many larvae No pupae  40000  tt  Control  25°  0  Jan .31" Feb.10  400  1  1  1  40*.  tt  5-6 days  29°  177 se^i? 5 3«^2$  0  1  Mar.11In heat Mar.17 Out Mar.24  192 89<y7<f  103 148  76 n  5 (2„5#) l o * 4?$ 0  No larvae 115 pupae - Sample of 15 Erne rge n t : 9 o-V>3 Larval : 3 10 p a r t l y emerged.  Table XXXVII - Heat Treatments (Continued) Cross; No.  400*  Temp. No. of Age When C. pi Treated Emerged  8 Xple orange 9?  4-5 days 29°  0  0  Mar.12In heat Mar.17 Out Mar.24  0  0  No larvae 108 pupae - sanple of 15 Emergent: 6 0 ^ 3 9 0 Larval : 6  3-4 days  29°  0  0  Mar.13In heat Mar.17 Out Mar.24  0  0  No larvae 126 pupae - sample of 15 Emergent: 3crkP299 Larval : 10  Control  25°  165  0  Mar .14** Mar.24  135 67 Co*  P Synpalpi 1  10 Xple orange <rV 4000  40000 A 1  1  p2 Synpalpi  Constitution.  »t,  it  830*829-  Life No. of History p2  6899  3 (2%) lo* 299  D i s s e c t i o n of pl- phpae  -  (iii)  5  7  -  Discussion;  In conjunction with both heat and cold treatments of Xple stock, w i l d stocks were also subjected to s i m i l a r temperatures. A l l f l i e s emerging from these treatments were normal - the F being normal.  1  and F  2  counts  This indicates that the Xple stock i s extremely sensitive  to variations from the normal temperature o f 25° C - since 15° C. r e duced the number of synpalpi appearing and was l e t h a l i n some age groups, and since 29° C. and 35° C. were both l e t h a l temperatures f o r the Xple stock.  (d) Investigation of the l e t h a l i t y of the mutant The p o s s i b i l i t y of a lethal character being associated with the mutant, synpalpi, was investigated i n two ways. The f i r s t method was to examine cultures of Xple and Xple synpalpi for l e t h a l i t y before emergence and the second was to test with C1B stock. This investigation was carried out since the preliminary crosses suggested a l e t h a l i t y and since Zaliss (1939) described a deficiency affecting the X chromosome whioh caused l e t h a l i t y i n the young larvae and eggs. (1) Crosses of Xple and Xple synpalpi. On A p r 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 ? ?  C  - 4 Xple orange synpalpi?? x 3 Xple orange synpalpi <rV  D -  x 3 Xple o"o»  4 Xple orange??  x 3 Xple orange^c"  1  On A p r i l 2 at 5 p.m. the parents were transferred to new bottles: A to A l , B to B 1, C to C 1, and D to II1. On A p r i l 3 at 5 p.m. another transfer of parents was made to new bottles:  A  1  to A , B 2  1  to B , C 2  1  to 0  and D  2  o  2  1  The parents were removed from A , B , 0  to D . 2  2  and D  2  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 Xple syno*  (emerged A p r i l 10: 21cf>32? - no synpalpi (died i n pupa case: 4 - 1? , 1°^ , 2 l a r v a l (^ .dead : 6%  - 59 A  : Xple syn? x Xple syno^  emerged A p r i l 11 : 29 o^M? died i n pupa case: 2 - l f f l , $ dead : 5$  A  : Xple syn? x Xple syn 0-7  control: emerged A p r i l -12 : 48 0^50$ - no synpalpi  B  : Xple? x Xple °^  emerged A p r i l 13 : 66o^ 63? -  2?, 1 or> died i n pupa case: 3-1 Or* 1? , 1 l a r v a l fa synpalpi: 2.3$ fa dead • :2.3$ fa  B : Xple? x Xple  emerged A p r i l 12 : 68 CT 66$ died i n pupa case: 3-2 o» 1? $ dead : 3$  B  ; Xple? x  control: emerged A p r i l 12 : 430* 39 ¥ -  C  t Xple orange syn. ? x Xple orange syn • cr>  emerged A p r i l 10 : 48 o 48? died i n pupa case: 2 - ley, 1 ? $ dead : 2$  A  Xpleo-7  1  71  C* : Xple orange syn ? x Xple orange syn cr?  emerged A p r i l 11 died i n pupa case $ dead  G  control: emerged A p r i l 12 : 2 3 ^ 35? - no synpalpi  2  : Xple orange syn.? x Xple orange syn. 0*  240" 43? - no synpalpi  D  : Xple orange ?x emerged A p r i l 11 : 52 54? - synpalpi: 3 ? Xple orange died i n pupa case: 16: 8or> 4 ? ) 4 l a r v a l fa dead : 15$ $ synpalpi: 2$  D  : Xple orange?x emerged A p r i l 11 : 470-T 63? - synpalpi: 1 ? Xple orange °^ died i n pupa case: 8: l < r » 4?, 3 l a r v a l fa dead : 7$ fa synpalpi: .9$  D  s Xple orange?x control: Xple orange emerged A p r i l 12 : 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 i n 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 number dead : 0  73?  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 i n 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 i n the Xple orange cross  (average 11.5$) and i n the Xple synpalpi cross (average 6.5$). In the d i s s e c t i o n 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  t h e i r e f f o r t s to free themselves, they contracted and expanded t h e i r bodies, forcing the front of the head out i n the region of the antennae. As mentioned before t h i s f o r c i n g 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 e f f o r t s may account f o r the squeezed appearance of the abdomen of some o f the synpalpi f l i e s which managed to extricate themselves.  (2)  C1B method.  The C1B method of detection of l e t h a l mutations i 3 limited i n that only sex chromosomes are involved i n the t e s t .  The GIB female used  i n the test has on one of i t s X chromosomes three factors on which t h i s method i s baseds  a known l e t h a l gene, 1, which k i l l s the males that  receive t h i s chromosome, sinoe these receive no normal a l l e l e i n 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 i t 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 males. 1  Such a cross  produces Bar and not-Bar females, half the males die because of the C1B lethal, and i f 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 males. 2  On February 21 the following crosses were set up: used were virgin). A  1 ClB/dl 49 ? x 1 Xple orange synpalpi cr  B  1 ClB/dl 49? x 1 Xple orange °^  C  1 lz/ClB  D  1 lz/ClB  E  1 lz/ClB  x 1 Xple synpalpi cr>  F  1 lz/ClB  x 1 Xple o"  0  1 lz/ClB 9.  x 1 Xple synpalpi cr  H  1 lz/CiB ?  x 1 Xple &  I  Control:  1  x 1 Xple orange synpalpi 0-^ ¥  x 1 Xple orange ^  1  7  1 lz/ClB ?- x 1 wildo-  7  The F counts are listed in Table XXXVIII A  (all females  -  63 -  Table XXXYIII A C1B ?¥ H ?? H (TV F  B  C  D  E  P  0  H  I  45 26 35 36 14 41 26 19 26 28 21 40 32 15 41 29 18 25 29 30 35 28 6 34 17 16 28  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?? A B C D E F a  H I  128 138 109 123 141 124 148 86 133  134 132 125 148 140 128 148 82 134  Synpalpi % Syn. F* orcf}  120 114 131 126 140 115 145 58 126  5 12 4 5 0 9 0 0 0  1.3 3 1.1 1.2 0 2.4 0 0 0  In the above table N ¥- and No- refer to those flies whioh re7  ceived the homologous X chromosome (not C1B) from the C1B mother. Ho lethal appears to be associated with the first chromosome v  sinoe the ratios were as expected:  1 C1B?- : 1 N ? : l o , Synpalpi 7  did appear in some of the males - this seems to indicate that i t i s 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 associated with Chromosome I alone. This method is extremely limited since it tests only Chromosome I; the lethality, therefore, may he associated 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  oounts are listed in Table XL.  1  Table XL  Feb.24 25 26 2? 28 Mar. 3  Wild Ye!Llow Hon ml Synpalpi Hon nal Synpjilpi ?? o-V ?? o V 99 crV 5 24 15 8 13 7 5 19 7 3 13 4 4 13 1 5 4 6  Total  66 40  The following 1 Y : 2 Y : Y : Y : 3  F  0  0  0 30  0  crosses were set up on February 25.  3 F yellowest 3 F yellow o-V 1  1  3 F virgin wild?* x 1 Xple orange synpalpi C  7  3 F virgin yellow?? x 1 Xple orange synpalpicr> 3 F virgin wild¥?-x 2 F yellow aV . 1  1  4  The F results are listed in Table XII. 8  6  7  -66Table XLI Yellow . Kor mal Synpalpi 9$ o-V 99  Y Mar.12 53 40 14 25 36 Total 78 76 1  Y Mar*12 22 14 6 26 Total  Nor mal 99  0 0  0  Synijalpi 9?  0  0  0  2  0 0  16 41 6 10 22 51  2  3 3 6  0 0  46 28 0 74  0  0  0 0  10 23 16 12 26 35  2 2  0 0  2  Y Mar*12 14 Total Y* Mar*12 14 Total  0  3  41 27 68  0  28 12 10 12 36 24  Expected resultst t  Y - a oross of F yellow ? x F yellow cr - since yellow 1  1  1  7  appears only in a homozygous oondition: the F results should he: 2  yellow or and yellow^ in a 1:1 ratio* 1  Y - a cross of F wild? x Xple orange synpalpi cf - since 2  1  1  the F wild? will have one homologue from the yellow parent and the 1  other from the Xple synpalpi parent, the F results should he: 2  2 Xple9 t  1 Xpletf" : 1 yellower . 1  7  Y - a cross of F yellow? x Xple orange synpalpi cr - since 3  1  7  the F* yellow ¥ is homozygous for yellow, the F^ results should he: 1 Xple 9 i l yellow cr . Y^ - a oross of F wild? x F yellowo^- sinoe the F wild$ will have one homologue from the yellow parent and the other from the Xple 7  1  1  1  synpalpi parent, the F results should he: 1 Xple* :1 yellow? :1 Xple©*: 1 yellower^ 2  67 * It was believed that the yellow synpalpi female had been prefertilized by a yellow male, sinee the cross of yellow?, x Xple orange cr  7  gives only yellow males and Xple females, whereas the actual F count 1  included yellow males and females as well as wild females. This assumption was further substantiated by the F results. 2  The actual F results conformed to the above expected ratios. 2 4 2  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 the F oonsisted of Xple males 2  2  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 i t , 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 Y the F con4  2  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 i t 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 D /ln (3L) PMe - Star located on 3  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 virgin curly dichaete? 1  x F curly diohaete cr , 1  7  The F results are listed in Table XLII. 2  Table XLII  Mar*14 17 18 19 Total  Curly dichaete Curly Dichaete &> a* cV n ?? 22 26 6 9 11 9 20 32 19 9 13 14 5 8 7 4 10 5 8 9 4 5 2 1 55 75 36 27 36 29  o-V 6 3 1 2 12  ? Xple 9% 7 9 3 2 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 or F progeny* I f the mutation 1  2  is on Chromosome II i t 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 i t would  >  -.69 have appeared with Curly since hoth homologues of Chromosome I I I were obtained from the mutant, or i n the last individual, ? Xple, as mentioned before.  The only significant result was that the r a t i o of males to females  i n the ? Xple individuals was; 2 females: 1 male, {iii)  Chromosome I I  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 v i r g i n ? x 1 Xple orange synpalpi<r>  B  1 Lobe Curly v i r g i n ? x 1 Xple orange  C  1 Lobe Curly v i r g i n ? x 1 Xple synpalpi o-">  D  1 Lobe Curly v i r g i n ? x 1 Xple &  E  1 Lobe Curly v i r g i n ? x 1 w i l d " 0  7  - control.  The F results are l i s t e d in. Table XLIII. Table XLIII i Lobe oV 27 ?? 35 Curly o-v 36 ?? 38  B  C  D  34 41 32 36  34 37 40 32  38 28 25 24  E (Control) 45 45 41 42  The F counts were as expected - 1 Lobe ? :1 Curly? :1 Lobe©'"': 1  1 Curly cr* . F  1  interses were set up consisting of a cross of 1 F v i r g i n 1  Curly ? x 1 F curly 1  .  The F results are l i s t e d i n Table XLI7. 2  4  -70 -  Curly  n A B C D E (Control)  154 137 150 165 139  Table XLIV Synpalpi  143 123 128 161 151  1  cr 2 crV 4 o-V 1 cr?  cr'o-  1  7  7  0  92 69 58 33 72  Synpalpi  7  '76 56 73 34 71  1? 0 1 cr? 0 0  ? U refers to the individual whioh has obtained homologues of the 2nd ohromosome from the mutant f l y . The expected -ratios i n 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 i n 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 i n Curly and US individuals - i n fact most appeared i n the Curly f l i e s . (iv)  Chromosome I I I  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 v i r g i n ?  x 1 Xple orange synpalpi or?  B - 1 Dichaete Glued v i r g i n ?  x 1 Xple orange cr  C - 1 Dichaete Glued v i r g i n ?  x 1 Xple synpalpio-  D - 1 Dichaete Glued v i r g i n ?  x 1 Xple ^  E - 1 Dichaete Glued v i r g i n ?  x 1 w i l d o - control.  The F  1  results are l i s t e d i n Table XLV.  7 7  7  7  - 71 Table XLV  Diohaete Glued  ? ? oV  ? $  The P  1  A  B  G  D  E (Control)  23 25  22 26  17 19  31 29  22 17  27 29  22 23  17 15  33 28  19 14  counts were as expected: 1 Dichaete? :1 Glued? : 1 Diohaeteo- :1 Glued 7  F  1  crosses were set up consisting of 1 P  1  v i r g i n Dichaete?  x 1 P Dichaete o1  7  2  F  results are l i s t e d i n Table XLVI. Table XLVI Dichaete Synpalpi cr'cfl ?? A B  C D E (Control)  108 120 125 128 176  176 127 189 196 191  0 0 0 1 cr? 0  Synpalpi 58 63 46 49 85  89 62 89 76 87  1 1 cr? 0 00 0  7E refers to the individual which has obtained hoth • homologu.es of the 3rd chromosome from the mutant f l y . The expected ratios i n 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 i n the ?N individuals since they had obtained both homologues of the 3rd chromosome from the mutant f l y . The F  2  results were very significant i n that the ratio of males  to females i n both Diohaete and ?U individuals was approximately  - 72 1 male:2 females - (normal r a t i o i s 1 : 1 ) . This may he explained by the fact that one h a l f the males received a Xple synpalpi or Xple X homologue from the original Xple synpalpi or Xple male - thus being homozygous for t h i s mutant character. A combination of t h i s 1 s t chromosome 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 h a l f the males to die.  - 73 -  (v) Summary Prom the preceding tests for linkage with Chromosomes I, II and III, i t 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 condition 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 absence or small number of synpalpi in the F  1  of~synpalpi crosses but its  reappearance in 2-8$ of the P progeny; and by the death of flies in 2  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; therefore, 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 i n the order named - yellow being at the very t i p .  The literature on a l l deficiencies  i n the region of yellow, aoheate)and scute was reviewed and many simi l a r i t i e s i n results to those recorded i n t h i s 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 phenotypic 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 i n the various preliminary crosses. Df(l)  260-1. Sight terminal bands are deficient proving lethal  i n 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 i n the Xple crosses of the present investigation. Q  Df(l)  sc * Paulson (1940),. The deficienoy whioh i3 i n 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 i n 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 i n the egg stage. Df(l)  260-2. The literature on t h i s 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 l e t h a l .  The females with the deficiency '  die i n the late pupal stage and the males i n 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 l a r v a l  period i s considerably lengthened i n Df(l) 260-2 as i t i s i n the case of synpalpi crosses. Df(l)  260-2, recorded by Sutton (1943), i s lethal i n 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 v a r i a b i l i t y i n 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 t o t a l mortality of this stock i s 33.4% death occurring at egg, l a r v a l , and pupal stages: 20.6 - 31.1$ as eggs. This high percentage of death i n the egg stage would account not only for the low counts from synpalpi crosses but also for the absence of dead larvae i n the examined cultures. The p o s s i b i l i t y that from 3 - Bfo of the deficient eggs hatch may indicate that the synpalpi, which appear i n t h i s 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 a i r 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 decreased 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 i n 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 D f ( 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 i n 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 i n the males which were hemizygous for the X chromosome but not i n the females whioh were heterozygous.  This  again indicates the significance of Chromosome I i n the inheritance pattern of the mutant. The absence of l e t h a l i t y i n the C1B test crosses, where brother s i s t e r 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  t h i r d chromosome, suggests  that Chromosome I I I i s probably associated i n some way with the production of the synpalpi mutation. The mutation i s not a simple deficiency of the X chromosome.  - 78 -  7  1.  Summary  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 o f the X chromosome alone,  but appears to involve the t h i r d Chromosome as w e l l .  r- 79 -  Literature Cited Braun, Weiner 1940,  Experimental evidence on the production of the mutant w  aristapedial 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 aristapedial M  1944.  Genetics, 27:173.  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 forehead and the fusion of 1  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 .  Fig. 8  Light source ventral.  Head of a synpalpi Xple fly, ventral view, similar to Fig. 6 . Light source ventral.  -  88  Fig. 8  - 89 -  Plate 7  Fig.  9  Head of a normal Xple f l y , l a t e r a l view, showing the position of normal antennae. Light source dorsal*  Fig* 10  Head of a synpalpi Xple f l y , l a t e r a l 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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