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J- INHERITANCE STUDIES OF WHITE-CAPPING IN YELLOW DENT MAIZE INHERITANCL STUDIES OF WSITE-CAPPING IN YELLOW DENT MAIZE Thesis Respectfully submitted in partial fulfillment for the degree of Master of Science at Michigan State College of Agriculture and Applied Science Chandrakant G. Kulkarni m 1926 THESIS - TABLE OF CONTENTS - Page. I--INTRODUOTION --------------------- 1 (1). Statement as to how the question arose- - l (2). Problem ----------------- l (3). Previous work on yellow endosperm color — 2 II--MATERIAL USED IN THIS INVESTIGATION ---------- 2 III'-TECHNIC ----------------------- 3 (1). Laboratory ---------------- 3 Classification and sorting ------ 3 (3). Field ------------------- 4 IV--DESCRIPTION OF THE WHITE CAP ------------- e ‘ V-—INHERITANOE OF WHITE CAP (Wcl) ------------ 6 (1). In Clement's White Cap ---------- 7 (2). In Folk's White Cap ----------- 7 (3). In Silver King - - ----------- 7 VI--1925 DATA FOR THE DETERMINATION OF THE DOMINANCE OF THE WHITE CAP -------------------- 8 VII--INHERITANCE OF WHITE CAPPING IN NORTHWESTERN DENT CORN - - - -_ --------------------- e TIII--INHERITANCE OF WHITE CAP (WOZ) ------------ 9 In Clement's White Cap ---------- 9 In Folk's White Cap ----------- 9' In Silver King ------------- 9 Page. IX--INHERITANCE OF WHITE CAP (W03) ------------- 9 In Clement's White Cap ----------- 9 In Folk's White Cap ------------- 9 In Silver King --------------- 9 X--SUMMARY ------------------------ 10 LITERATURE CITED --------------------- ll ACKNOWLEDGMENT ---------------------- 12 TABLES, 1-25 ......... - ------------- 13 - 1 - (1). Statement as to how the question arose. The late PrOfessor F. A. Spragg had observed from time to time in the course of his researches and observations on corn that when a yellow corn is pollinated by some white corns the resulting g kernels of these crosses were white-capped thus indicating that white-capping was dominant to yellow-capping. He would have investigated the occurance of the white cap, but due to the plant breeding problems of a more important nature which he had at hand, he did not go on with it further, and the problem was allowed to rest. A new impetus was given to the problem of investigating white cap when Mr. J. R. Duncan, Research Assistant at the Michigan Agricultural Experiment Station brought for analysis to PrOfessor F. A. Spragg a few ears of corn some grains of which were white-capped. These kernels were the open pollinated crosses between Duncan and Silver King, Duncan being the female parent. The former is a yellow corn and the latter a white corn. (8). The problem. As stated above, in the case of dent corns, white cap was observed to be dominant to the yellow cap although no exact investigation had been carried on up to 1924, when the writer came to work under the late Professor F. A. Spragg as a graduate student. All the material pertaining to the white cap was given to the writer for further investigation. The problems to be investigated were:- (1) Whether white cap was due to a Mendelian factor or purely to the effect of the environmental conditions. (2). Is this factor an inhibitor, and if so, what tissues does it affect? (3) Previous work on the yellow endosperm color. An observation of a white-capped kernel reveals that the yellow pigment, so characteristic of a yellow corn, is entirely lacking in the cap region at the crown. This shows that whiter capping is a condition of the endosperm. Therefore it was thought desirable to review very briefly the previous work on the yellow endosperm color. Two factors for the yellow endosperm color in maize have been described. The factor (Y) was first described by East (1910) although Correns (1901) had described it without giving it any symbol. Emerson (1911) also describes it. The factor (Tp) was first described by East (1910) and Emerson (1911). Correns (1901) describes a pale aleurone pigment in maize and this pale yellow was shown by Kvakan (1924) to be identical with brown aleurone Gan). (The factor (Y) gives a dark yellow color, when the modifying factors are present the factor (Y) also gives a deep orange color in the endosperm (Anderson 1924). This orange color is due to the (Y) factor together with the modifying factors because the kernels having the orange color show linkage with (P1) factor, a factor for plant color in maize (Emerson, 1921; Anderson, 1924.). ‘II--Material used in this investigation. As all the material used in this investigation came from different sources, it was thought desirable to tabulate it to make it more intelligible to the reader. The following material was used in this investigation. Accession number Name Description 235 Duncan Yellow-cap yellow 254 Duncan Yellow-cap yellow 257 Bailey Connecticut Yellow-cap yellow yellow 265 Clement's White Cap White cap yellow 266 Folk's White Cap White cap yellow 26? Silver King White 286 Northwestern Dent Red Pericarp Accessions 235 and 254 are two inbred strains of yellow- cap yellow endosperm corn. Accession 257 is a cross between two inbred yellow-cap yellow endosperm strains. Accession 265 and 266 are two different strains of white cap yellow grown in the State of Michigan. Accession 267 is a commercial variety cf the white corn. Accession 286 is also a commercial variety of corn. Accessions 235 and 254 were received from the Farm CrOps department of Michigan State College. Accession 257 was given to the writer by the late Professor F. A. Spragg, while Accessions 265, 266, 267, and 286 were re- ceived from Mr. J. R. Duncan, Research Assistant at the Michigan Agricultural Experiment Station. I III—-Technic. (1). Laboratory. Classification and sorting. For the purpose of this investigation the above material -4- was sorted into four classes: (1). Those grains whose caps were Judged to be as white as any caps on the established white-capped variety, furnished by Mr. Duncan. . (2). Grains with white caps all somewhat darker than the light- est, some of them coming from the ears also showing segregation. (3). This class consisted of grains whose caps were still darker than those in the second class. These came from the ears which were segregating for the pure yellow-capped grains. (4). Ordinary yellow-capped grains. For the purpose of classification there were assumed three dominant factors (W01, W03, W03.) and their allelomorphs, which are considered to be the absence cf white capping. Wcl is considered to be powerful enough to produce a fully white-capped yellow in the presence cf (Y) the factor for the yellow endosperm. W02 is considered less powerful, and We still less power- 3 fun producing only light cap of very slight intensity. This hypothesis was not in the end substantiated, but was found useful in the analysis Of the data. It worked out there was only one factor for the white capping, the other conditions being due to modifying factors. It was comparatively easy to sort the material in 1923 as the material received was well matured. But in 1924, due to the shortness cf the growing season, classification was difficult and the material was not entirely mature when harvested. If the -5- material is even slightly immature it is very hard to differentiate the various shades cf white—capoing. In 1925 the material was in an excellent condition for classification, as it had a long grow- ing season and hence was thoroughly mature. (2). Field. The following method cf selfing and crossing was used:- Various methOds cf corn pollination have been tried and some of them have been found to be satisfactory and the others are still in the process of experimentation. As soon as the ear buds appeared they were sacked with the glacine bags before any silks appeared. These bags were fastened in such a manner with a clasp that the wind would not blow them away and at the same time the ear buds would have ample room for their develOpment.~ The ear buds were watched from day to day. (If two or three inches of silks are exposed, about a third of the kernels are represented, when six inches cf silks are exposed at least three-fourths of the kernels are represented). When the silks were six inches long pollination was made. The tassels were sacked the day prior to the pollination, so that abundant viable pollen might be obtained for the pollination the next day. In sacking the tassel twelve-pound grocer's paper sacks Of heavy quality (NO. 1., 40 1b. Kraft, pinch bottom type) were used. The heavy grade cf paper had the advantage of not being torn away by the wind. The two upper leaves were removed before the tassel was sacked. The leaves were not sacked because they transpire and in- Jure the pollen and sometimes collect large quantities of pollen on their surface. The sacks were clasped firmly with a clasp around the stalk below the tassel to prevent the tassel from being broken by the weight of the sack. To accomplish this in a most satisfactory manner, the mouth Of the sack was folded as it was folded in the -5- bundle, and then the corners were folded towards the stalk and the folds, thus formed, were turned together and fastened with a clasp. This method would also keep the sack from being blown away in a heavy wind. The tassel sack was then dated. The pollination was done in the following way:- The tassel sack was shaken horizontally and unfastened very carefully to prevent the loss Of any pollen. The anthers and the other rubbish that first appear were discarded while the main mass of the pollen still remained in the sack as the fine yellow powder. The glacine sack on the ear was torn off at the top to make an Opening and the contents cf the pollen sack were carefully emptied onto the silks. The glacine sack was then given a light shake to affect an abundant pollination. The pollen sack was used to cover the ear and fastened with a clasp in such a way that it would not be blown away by the wind, and thus eliminate the danger of foreign pollination. A second date, the date of the pollination, was written on the outside of the sack. A small tag with the date of the pollination, the nature of the pollination (crossed or selfed) and the pedigree number was tied, just above the ear, to the stalk that the ear might be recognized in case the sack was blown away by the wind later in the season. The sacks were left on the ears until the harvest time. In the above method no alcohol was used to disinfect the hands after each pollination, as many investi- gators suggest, as the hands do not come in contact with the silks. IV-Description of the white cap. White-capped grain is easily distinguished from an un- modified yellow cap by the fact that the apex 0f the grain is white. -7- Sections of the grain show that the soft starch portion of the endosperm which reaches the apex is in one case pure white and in the other slightly tinged with yellow. The yellow pigment that is so characteristic of the yellow grains having the yellow endosperm is entirely lacking the cap region and as one examines a kernel having a white cap the characteristic yellow pigment is seen clearly just below the cap. Hence, the white-capping is due to the elimination of the color by an inhibitor. V--Inheritance of white cap (Wei). (1). In Clement's White Cap. In order to determine the inheritance of Clement's White Cap it was crossed by Bailey Connecticut Yellow, the former being the female parent. These.orosses were made in the year 1924. The F1 seeds Of these crosses gave only grains that were white-capped yellow. In'Table l are summarized the results of these crosses. When the F3 white-capped seeds cf the above crosses were selfed in 1925, they gave a ratio of 3:1 of the white-capped grains to the yellow grains, thus establishing that the white-capping in this case was due to a single domirant Mendelian factor. Table 2 shows the results of the analysis of these crosses. That the white capping is due to a single factor in this strain can also be seen by the results of the backcrosses which ' are summarized in Table 3. (2). In Folk's White Cap. Folk's White Cap gives the same results as the Clement's White Cap. When it is crossed by a yellow the F1 kernels are all white- capped yellow (Table 4). When these F1 kernels were selfed they gave a ratio in F5 of three white-capped grains to one yellow-capped grain -3- (Table 5), showing that the white-capping in this strain is also due to a single dominant Mendelian factor. The backcrosses substantiate the same conclusions (Table 6). (3). In Silver King. In order to establish the existance of a white-capping factor definitely in Silver King, which might not be able to express it- self in the absence of the yellow, Silver King was crossed by Bailey Connecticut Yellow. The F1 seeds of these crosses were all white- capped yellow, thus proving the existence of a white capping factor in Silver King. In Table 7 are summarized the results of these crosses. These white-capped seeds of F1 generation when selfed the next year, gave a ratio of 3:1 for yellow to white grains. (Table 8) The yellow grains on further classification of the white caps gave a ratio cf 3:1 for the white caps to the yellows. (Table 9). The backcross data substantiate the same conclusions. (Tablelrn This indicates that the white cap obtained by crossing Silver King x Bailey Connecticut Yellow acts in the same Way as did the Clement's White Cap and the Folk's White Cap, in that it is dominant in F1 and segregates into a 3:1 ratio for the white-capped to the yellow-capped grains in F2. VI—-1925 data for the determination of the dominance of the white cap. The material was very hard to classify in 1924 due to the extreme immaturity and because some kernels on each ear could not be classified (Tables 1, 4, 7, last column) it seemed desirable to make some crosses in 1925 of white caps onto the yellow in order to determineziefinitely the dominance of the white cap. With this aim in view, crosses were made between Duncan and Clement's White Cap, Duncan and Folk's White Cap, and Duncan and Silver King. -9- The F1 data of the cross Duncan x Clement's White Cap, as summarized in Table 11, those between Duncan and Folk's White Cap, as summarized in Table 12, and those between Duncan and Silver King, as summarized in Table 13, indicate that the white- capping is dominant in all the cases, because all the grains from these crosses were only white-capped. VII-~Inheritance of white-capping in Northwestern Dent Corn. Northwestern Dent corn, with a colored pericarp except in the cap region, having been selected for the white endOsperm was crossed onto Duncan. The resulting Fl grains were all white- capped yellow (Table 14) thus proving that in this case also the white-capping factor was dominant. VIII--Inheritance of white cap (Wcz). The F data and the backcrosses both in Clement's White Cap, 2 Folk's White Cap and Silver King show We to be dominant to the yellow-cap and give a ratio of 3:1 for tie white-capped to the yellow-capped grains in the F2 generation, when selfed. The white caps in the F5 generation are Of all the shades (Wcl, W03, W03). This indicates that there must be modifying factors that cause the various shades of the white-capping. If there should‘be no modify- ing factors it would be expected that it would segregate in F5 in a distinctly 3:1 ratio for Wag to yellow on the basis of independent inheritance, which is not the case. (In Tables 15, 16, 17, and 17A) are listed the results of these crosses. The presence of the modifying factors can also be ascertained by studying the backcrosses ch x (WczY x'ch) from each of the above strains. The results of these backcrosses are listed.in Tables 18, 19, and 20). -10- If Wc8 was an independent factor it would segregate into a 1:1 ratio for ch to yellows in the backcrosses mentioned above, which is not the case. The white caps obtained are not only Wcz but all the shades (Wcl, Wcz, W03). This again substantiates the conclusions made above as to the modifying factors. IK-—Inheritance of white cap (W03). W03 from all the three strains mentioned above acts the same way as Woz in all the crosses and the backcrosses as can be seen from the data presented in Tables 21, 22, 23, 24, and 25. The data also indicate that Wc3 is not inherited independently as was assumed at the beginning, but is the outcome cf the action of the modifying factors. ‘ X--Summary. (1). A white-capping factor is described and is designated as We. It is inherited in a simple Mendelian way, the character be- ing due to a single factor. It is dominant to the yellow-capping in the first generation and segregates in the second generation in a 3:1 ratio for the white-capped grains to the yellow-capped grains in Clement's White Cap, Folk's White Cap, and Silver King. (2). White-capping is due to an inhibitor which drives the color away from the crown cf the kernel. (3). Silver King, a white corn, carries a white capping factor but is unable to express it in the absence of the yellow endOHperm. (4). Various shades cf white-capping are possibly due to the presence of modifying factors. -11.. *LITERATURE CITED- Anderson, E. G. 1925. Genetic factors for the yellow endosperm color in maize. Michigan Acad. Sci. Arts and Letters 4:51-54. Correns, C. 1901. Bastards zwischen Maisrassen, mit besonderer Berucksichtigung der Xenien. Bibliotheca Botanica 53:1-161. East, Edward M. 1910. A Mendelian interpretation of variation that is apparently continuous. Amer. Nat. 44:65-82. East, E. M. and Hayes, H. K. 1911. Inheritance in maize. Connecticut Agr. Exp. Sta. Bul. 167:1-162. Emerson, R. A. 1911. Latent colors in corn. American Breeders' Association Dept. 6:233-237. Emerson, R. A. 1921. The genetic relations of plant colors in maize. Cornell Univ. Agr. Exp. Sta. Memoir 39:1-156. Kvakan, Paul. 1924. Inheritance of brown aleurone in maize. Cornell Univ. Agr. Exp. Sta. Memoir 83:1-22. Spragg, F. A. Some unpublished records cf the Michigan Agr. Exp. Station. -12.. ACKNOWLEDGMENT It is very unfortunate to see that Professor F. A. Spragg who helped the author to lay out this piece cf research and gave many suggestions did not live to see the work completed. To his memory the author humbly dedicates this work, as this is the first piece of research turned out after his death by one of his students. The author is indebted to Professor E. E. Down and Mr. H. M. Brown for ad- vice and criticism thruout the work and the pre- paration cf the manuscript; to F. H. Clark for valuable suggestions. Appreciation is due to Professor J. F. Cox, Professor E. E. Down, Mr. H. M. Brown and Doctor E. A. Bessey for the final review cf the manuscript. Thanks are also due to Professor H. H. Bartlett and Doctor E. G. Anderson, University of Michigan, for reading over and making many valuable suggestions on the manuscript. The writer is thankful to Mr. J. R. Duncan, Research Assistant, at Michigan State College for supplying the material. -13- Table 1. F1 progeny of the cross Clement's White Cap (WOY) Ac. 265 x Bailey Connecticut Yellow (ch) Ac. 257. N0. 3 W01 : W03 : Wcs : Kernels that could not be classi- : : : fied due to immaturity. 48901 i 188 f 51 5 62 3 13 48902 i 215 E 55 g 54 E 0 48903 E 311 E 195 E 196 E 0 48904 E 295 g 70 g 118 E 0 48905 i 343 E 164 E 147 i 0 There were about twenty ears of this cross but only five of them were classified, the others were discarded due to ex- treme immaturity. No. 48905 was reserved for further planting. -14- Table 2. Fé pr0geny of the cross Clement's White Cap (WcY) Ac. 265 x Bailey Connecticut Yellow (ch) Ac. 257. N0. : White Cap Yellow : Yellow : Deviation from 3:1 3 (WclY) E (Y) 3 ratio. 544802 572 129 : 3.75 1 8.54 544804 289 94 1.75 1 5.72 544808 558 I 120 5.50 1 8.25 544808 A 317 109 2.50 1 8.05 544808 525 184 7.25 1 7.77 544809 298 100 1.00 1 5.81 544813 310 1:30 2.50 1 5.91 544815 145 53 4.00 1 4.09 544815 A g 430 g 141 g 1.75 i;6.98 544818 585 120 0.75 1 8.42 544819 512 108 1.50 1 5.97 544819 A 588 ' E 127 5.25 1 8.50 544821 187 61 1.00 1 4.60 544825 404 148 10.00 1 8.88 Total : 4652 E 1592 E Calculated : g 2 3:1 : 4683 : 1561 : Deviation 31.00 1 33.07 ‘— -15- Table 3. PrOgenies of the backcrosses Duncan (ch§ Ac. 355 x (Clement's White Cap (WcY) Ac. 265 x Bailey Connecticut Yellow (ch) Ac. 257) No. E White Cap Yellow E Yellow E : (WclY) : (Y) : 542917 1 544812 E 158 E 209 E 542902 x 544810 E 212 E 218 E 542917 1 544821 E 194 E 199 E 542920 x 544808 E 148 E 151 E Total E 712 E 775 E Calculated 1:1 E 745.5 E 745.5 E Deviation 31.5 1 13.01 -16- Table 4. F1 progeny of the cross Folk‘s White cap (WcY) Ac. 266 x Bailey Connecticut Yellow (ch) Ac. 257. N0. : W01 : We2 : W03 : Kernels that could not be : : : : classified due to immaturity. 49001 E 198 E 125 E 109 E 5 49002 E 505 E 70 E 77 E 18 49005 E 505 E 145 E 47 E 10 There were about thirty ears of this cross of which only three were classified the others were discarded due to extreme immaturity. No. 49002 was reserved for further planting. -17- Table 5. F2 progeny of the cross Folk's White Cap (WcY) A0. 288 x Bailey Connecticut Yellow (ch) Ac. 257. No. : White Cap Yellow : Yellow : Deviation from 3:1 : (WclY) : (Y) : ratio. 545101 E 554 E 125 E 5.75 1 8.58 545101 A E 284 E 91 E 2.75 1 5.88 545105 E 214 E 75 : 2.75 1 4.97 545105 A E 527 E 112 3 2.25 1 8.12 545108 E 575 E 125 E 1.00 1 8.50 545109 E 281 E 98 E 8.75 1 5.52 545110 E 361 3 128 E 5.75 1 8.48 545115 E 277 E 89 E 2.50 1 5.59 Total E 2451 E 857 E Calculated E E - 3:1 : Z466 : 882 : I Deviation ' 15.00 1 16.73 -13- Table 6. Progenies of the backcrosses Duncan (ch) Ac. 235 x (Folk's White Cap (WcY) Ac. 288 x Bailey Connecticut Yellow (on) Ae. 257A No. : White Cap Yellow : Yellow : : (WOIY) : : 542909 x 545107 E 198 E 198 E 542941 x 545112 E 172 E 187 E 545945 1 545115 E 280 E 254 E 542945 1 545117 E 180 E 180 E Total E 810 E 797 E Calculated 1:1 E 805.5 E 805.5 E Deviation 6.5 1; 13.52 -19- Table 7. F1 progeny 0f the crOss Silver King (wcy) Ac. 267 x Bailey Connecticut Yellow (ch) Ac. 257. No. We We Wc Kernels that could not be E 1 E 2 E 3 1f» classified due to immaturity. 49101 E 108 E 85 E 199 E 145 49102 E 59 E 57 E 81 E 0 49105 E 79 E 58 E 125 E 141 49104 -E 70 E 52 E 47 E 185 49105 E 128 E 58 E 91 E 258 There were about thirty five ears of this cross but only five were classified the others were discarded due to extreme immaturity. No. 49101 was reserved for further planting. -30- Table 8. F2 data or the Cross Silver King (wcy) Ac. 267 x Bailey Connecticut Yellow (ch) Ac. 257. No. : Yellow : White : fiEviation from 3:1 ratio ’_' f (Y) 3 (y) 3 545501 527 101 8.00 1 8.04 545501 A 502 185 E 1.75 1 7.54 545502 594 ~ 154 2.00 1 8.71 545505 204 82 - 4.50 1 4.78 545504 587 148 12.75 1 8.74 545509 197 E 84 1.25 1 4.72 545511 577 114 E 8.75 1 8.47 545515 497 174 8.25 1 7.57 545514 285 E 72 11.75 1 5.55 Total E 5148 E 1052 E Calculated E E E 3:1 : 3135 : 1045 : Deviation 13 .00 1 18 .86 -31- Table 9 0 Further classification or the yellow grains 0f Table 8. No. : White Cap Yellow : Yellow : Deviation from 3:1 3 (WclY) E (Y) E ratio. 545501 E 340 E 87 E 5.35 i 5.38 545501 A E 375 E 137 E 1.50 i 6.54 545502 291 105 E 4.50 1 5.80 545505 152 52 E 1.00 1 4.17 545504 292 95 1.75 1 5.75 545509 149 48 E 1.25 1 4.10 545511 282 95 E 0.75 1 5.87 545515 575 124 0.25 1 8.51 545514 198 E 85 0.75 1 4.74 Total E 2552 E 798 E Calculated E E E 3:1 : 2361 : 787 ° Deviation E 9.00 1_l6.38 -23- Table 10. Progenies of the backcross Duncan (ch) Ac. 254 x (Silver King (Woy) Ac. 267 x Bailey Connecticut Yellow (ch) Ac. 257. N0. E White Cap Yellow E Yellow E : (W01 ) : (Y) : 545914 1 545509 E 202 E 212 E 548052 x 545511 E 150 E 114 E 545808 x 545514 E 128 E 155 E Total E 480 E 481 E Calculated 1:1 E 480.5 E 460.5 E Deviation 0.5 1 8.86 Table 11. F progeny 0f the cross Duncan (ch) 1 White Cap (WcY) Ac. 265. Ac. 235 x Clement's No. E White Cap Yellow E Yellow : : (WcY) : (Y) : 542957 x 542512 E 555 E 0 E 542212 x 542517 E 488 E 0 E 542801 3 542518 E 455 E 0 E 542952 x 542402 E 587 E 0 E 542804 x 542405 E 552 E 0 E 542805 x 542408 E 499 E 5 E 542258 1 542418 E 585 E 0 E 542805 1 542505 E 452 E 0 E 542005 x 542505 E 525 E 0 E 842918 3 542507 E 404 E 0 E 542019 x 542515 E 525 E 0 E 542924 x 542802 E 287 E 0 E 542927 x 542808 E 588 E 0 E 542808 x 542812 E 284 E 0 E 542911 x 542708 E 523 , i o E 542902 x 542705 E 514 E 0 E 542905 x 542704 E 297 E 0 E 542920 x 542710 E 257 E 0 E -34- Table 12. F1 progeny or the cross Duncan (ch) Ac. 235 x Folk's White Cap (WcY) Ac. 288. N0. E White Cap Yellow E Yellow E : (WcY) E (Y) E 542921 x 545002 E 295 E 0 E 542915 x 545004 E 225 E 0 E 542808 x 545108 E 557 E 0 E 542811 x 545111 E 589 E 10 E 542818 x 545112 E 402 E 0 E 542942 x 545502 E 459 E 0 E 545702 x 545508 E 298 E 0 E 542958 1 545512 E 508 E 0 E 542955 1 545515 E 244 E 0 E 542817 x 545518 E 158 E 0 E -35- Table 13. F1 pr0geny of the cross Duncan (ch) Ac. 235 1 Silver King (ch) Ac. 267. No. E White Cap Yellow E Yellow E : (WcY) E (Y) E 542951 1 545408 E 578 E 0 E 542218 3 543408 E 482 E 0 E 542814 x 543409 E 256 E 0 E 542234 1 543415 E 247 E 0 E 543728 x 543601 E 308 E 0 E 543801 x 543804 E 92 E 0 E 542933 x 543806 E 358 E 0 E F progeny or the 1 cross Duncan (on) Ac. 235 x Northwestern Dent (ch) Ac. 286. -26- Table 14. : White Cap Yellow : Yellow : : (WcY) : (Y) : 542223 544501 : 436 3 0 : 546010 544503 i 368 g 0 : 544705 544507 : 289 i O : 542219 544508 : 536 i O i 546009 544511 : 395 i 0 : 542237 544513 : 145 : O : 542226 544514 : 375 4 g .0 00 00 00 co Table 15 e F: progeny of the cross Clement's White Cap (WczY) A0. 265 x bailey Connecticut Yellow (ch) A0. 25?. W02 grains from F l planted and selfed. N0. E WolY WczY E W03Y E Total white yelloszeviation from : : : : cap yellow : : 3:1 ratio 544901 E 55 E 505 E 155 E 525 E 175 E 2.25 _-_+_ 7.74 544902 E 79 E 292 E 124 E 495 E 150 E 5.75 i 7.47 544904 E 73 E 255 E 129 E 457 E 170 E 5.75 1 7.49 544905 E 55 E 149 E 95 E 500 E 97 E 2.25 1 5.52 544905 E 54 E 257 E 93 E = 424 E 141 E 0.25 i 5.94 Total E 555 E1299 E 574 : 2251 E 745 E Calculated: E 2252.75 E744.25E 3:1 : : : : Deviation 1.75 ;I_- 15.94 A Table 16. F2 progeny or the cross Folk'a White Cap (WcY) Ac. 266 x Bailey Connecticut Yellow (ch) A0. 25?. W0 2 grains from F1 planted and selfed. No. E WclY E WcaY E W03Y E Total white E yellow E Deviation from : : : : cap_yellow : : 3:1 ratio 545202 E 45 E 108 E 55 E 220 E 74 E 0.50 1 5.01 545204 E 60 E 143 E 100 E 303 E 109 E 6.00 1 5.93 545208 E 52 E 124 E 83 E 259 E 93 E 5.00 i 5.48 545209 E 30 E 77 E 55 E 162 E 60 E 4.50 £14035 545211 E 43 E 105 E 67 E 215 E 76 E 3.25 ¢;4.98 Total E 231 E 557 E 371 E 1159 E 412 E CalculatE E 1178.25 E 392.75 E ed 3:1: : : : * Deviation 19.25 1 11.58 Table 17. F3 progeny of the cross Silver King (ch) A0. 267 x Bailey Connecticut Yellow (ch) Ac. 257. W02 grains planted and self pollinated. N0. E Yellow E White E Deviation from 3:1 ratioE _ 3 (Y) 3 (Y) 3 : 545505 E 384 E 128 E 0 i 5.51 E 545505 : 589 E 115 E 10.25 :t 5.55 E 545508 E 545 E 99 E 12.00 i 5.15 E 545510 E 585 E 122 E 5.00 i 6.58 E Total E 1504 ' E 455 E E x : : : ___ CalculatedE E E E 551 : 1475.75 492.25: : Deviation 27.25 1 12.96 -30.. Table 17 A. Further classification of the yellow grains of Table 17. No. E WclY E Wc Y E We Y E Total whiteE YellowE Deviation from : : 3 : 3 : cap yellow : ELY) : 3:1 ratio 545 : : : 63 : 289 : 95 : 1.00 1;5.72 545505 E 45 E 179 E 55 E 290 E 99 E 1.75 1:5-76 545508 E 55 E 170 E 55 E 259 E 85 E 0.25 i 5.42 545510 E 49 E 157 E 74 E 290 E 95 E 0.50 1 5.74 15551 E 171 E 701 : 255 E 1125 E 575 E Calculat E E E E 1128 E 575 E __ed 3:1: : ° : : : Deviation 0 1 11.33 -31- Tabl. 180 Progeniee or the backcross Duncan (ch) Ac. 254 x (Clement's White Cap (IczY) A0. 265 x Bailey Connecticut Yellow (ch) Ac.257) W03 grains planted. No. W03Y Total white Yellow cap yellow ‘ (Y) 00 .0 O. I: 0 H K: 00 co co 2: 0 N 0-4 00 00 O. O. .0 I. O. O. .0 543914 x 544903 FJ ...: (I) 83 229 224 . .. .. N (0 545914 x 544909 m 0 to i:- 45 159 198 542906 I 544910 : 61 204 1 [0 IF 0. 00 00 O 00 O o 00 00 no ._. . F; . (A o. O 00 o o O o no 0 00 Total ‘2 03 OJ N U! . 188 586 600 o. o. o. 00 00 o. o. o. 00 00 Calculated 1:1 593 593 to co oo oo o. co .0 o. oo oo 00 oo 00 Deviation 7.00 1 11.60 -33- Table 19. PrOgenies or the backcross Duncan (ch) A0. 254 x (Folk'e White Cap (WcY) Ac. 266 x Bailey Connecticut Yellow (on) Ac.257). Wc grains planted. 2 No. E WclY E We Y E Wc Y E Total White E Yellow : : 2 : 3 : Cap yellow : 545905 1 545202 E 22 E 55 E 59 E 124 E 110 546028 x 545205 E 54 E 89 E 54 E 177 E 155 544711 x 545211 E 44 E 107 E 55 E 215 E 249 75551 E 100 E 259 E 158 E 517 E 525 Calculated; 1:1 E E 521 E 521 (I! (1) Deviation 3. E3 l+ H C) -33- Table 20. Progenies of the backcross Duncan (ch) A0. 254 1 (Silver King (Woy) Ac. 267 x Bailey Connecticut yellow (ch) A0. 25?. W02 grains planted. No. : WclY : WczY : W0 Y : Total white : Yellow : : : 3 : cap yellow. : (Y) 544721 x 545502 E 9 E 47 E 15 E 72 E 75 545014 x 545505 E 15 E 77 E 50 E 122 E 154 Total E 24 E 124 E 45 E 194 : 207 Calculated 1:1 E E 200.5 . 200.5 Deviation 6.5 _-I_- 6.75 -34- Table 21. F2 progeny of the cross Clement's white cap (Wch) Ac. 265 x Bailey Connecticut yellow (ch) Ac. 257. W03 grains from F1‘ planted and selfed. No. E WclY E WozY E Wch E Total white EYellow E Deviation 3 = 3 3 cap yellow : (Y) : from 3:1 : E 3 E s E ratio. 545001 15 E 25 E 48 E 85 E 29 E 0.25 i 5.15 545004 E 44 E 105 E 300 E 349 E 118 E 1.25 j;6.31 545005 51 85 125 242 75 4.25 1 5.20 545011 55 84 155 274 91 0.25 i 5.58 545012 54 129 248 451 141 2.00 .t 5.99 545014 E 34 E 88 E 180 E 303 E 98 E 2.00 i 5.84 Total E 213 E 515 E 956 E 1684 E 552 E Calculat E E 1577 E 559 E eg 3:1: Deviation 7.00 1 15.85 ‘ -35- Table 82. ‘ F2 progeny of the cross Folk'a white cap (Wch) Ac. 885 x Bailey Connecticut yellow (ch) A0. 257. We3 grains from F1 planted and selfed. No. E WclY E WozY E W03Y E Total white E Yellow E Deviation : : : : cap yellow : (Y) : from 3:1 : E j, : 1 : ragig;____ 545501 E 59 45 149 254 85 E 1.25 i 5.55 545302 E 89 E 65 E 232 E 385 E 127 E 1.25 i 6.62 545305 E 101 E 79 E 250 E 440 E 150 El0.00 i.7-15 545307 E 73 E 55 E 178 E 305 E 97 E 3.50 ¢_5.86 545308 E 97 E 74 E 238 E 409 E 139 E 3.00 i_6.84 545510 107 81 250 448 145 5.25 :t 7.11 Total E 525 E 400 E1317 E 8242 E 751 E Calculat E E .2244.75 E 748.25 E __ed 3:1: : : : Deviation 2.75 1 15.97 -36- Table 23. Pr0geniee of the backcross. Duncan (ch) A0. 254 x (Clement's white cap (W03Y) Ac. 255 x bailey Connecticut yellow (ch) Ac.257) W03 grains planted. No. : WclY 2 We Y 3 W0 Y E Total white E Yellow 2 : 3 : 3 :_ cap yellow : (Y) 545807 x 545004 E 19 E 42 E 77 E 158 E 141 545009 x 545010 E 15 E 55 E 57 E 118 E 119 542915 x 545011 E 29 E 59 E 128 E 225 E 250 Total E 55 E 147 E 272 E 482 E 490 Calculated 1:1 E E 486 E 486 Deviation " 4 .00 i 10 .51 -37- Table 24. Progeniee or the back cross Duncan (ch) A0. 254 x (Folk'e white cap (Wes!) A0. 265 x Bailey Connecticut yellow (ch) A0. 25?.) W03 grains planted. N0. : WclY 3 WozY 3 W0 Y 1 Total white 3 Yellow 3 ~ 3 3 3 3 cap yellow 3 (Y) 544707 1 545504 E 29 E 22 E 57 E 118 E 128 544504 x 545508 ~E 50 E 58 E 118 E 205 E 208 Total E 79 E 50 E 185 E 524 : 555 Calculated 1:1 E E 550 E 550 Deviation 5.00 .t 8.55 -38- Table 25. Progeniee of the backcross Duncan (ch) A0. 854 x (Silver King (ch) A0. 257 x Bailey Connecticut yellow (ch) A0. 25?.) W03 grains planted. No. E WclY : W0 Y E We Y E Total white E Yellow : : 3 : 3 : cap yellow : (Y) 545024 1 545704 E 21 E 50 E 78 E 129 E 150 545905 x 545710 E 17 E 22 E 71 E 110 E 115 545904 x 545715 E 25 E 55 E 59 E 128 E 155 Total E 51 E 88 E 218 E 557 : 578 Calculated 1:1 : E 572.5 : 572.5 Deviation 551920 'MLER-s.c_t_1___AIRER co_. 511111055113.htm“; BINDERS ‘ Omce omrmens ANN ARBORMICHIGAN