_ ., ,._(u.u...:mm; .m 'n-mhm‘Iurul‘m-ql’, u zuum Emu-1.: uni-1‘. mkmwowew-l “mung-Una I ' M up n. ma- 1 r" at”. '-9‘l\-¢M/uak» um ~ MHH-AU—"LY‘.JIII” y '11 .a.’ \ , tvC‘IL4] ‘11 ’ (/LC 1’. 1;. fl ._.. H 15*- .- 3:9“.‘3’13...’ iSC‘Di‘iii‘w r'IIalfiil. aft-1. ~|pll|| ill [III] I II! .n 4“" BREEDING OF SEBAIKS OE A‘s-TESTER YELLO‘A' DJZIT CORN Thesis Respectfully auhnitted in partial fulfillment for the Degree of Doctor of PhiIOSOPhy at Michigan State Co 11636 of Agriculture and Applied Science -0- Kuo-J-Iing Liu 1927 THLbIS -AC£€O'.«1&JM" "14'“— The writer wishes to aoimoxaledge his indebtedness to Professor E. E. Down and Mr. H. LI. Brown for their generosity in transferring him the material of the experiment which had been already under way, and their valuable suggestions and thorough criticisms of the following thesis. He is indebted to Mr. F. H. Clark for very helpful suggestions on the manuscript. He is also indebted to Lirs. liurian Brown for corrections of English in the manuscript. Appreciation is due Professor J. E. Cox, Dr. E. A. Bessey, and yr. 11. P. Hibbard for the final review of the thesis. 10:32:31- TABLE OF CONTENTS Rise INTRODUCTION --------------------------- 1 MATERIALS.KRDI ETHOLD UgED .................... 5 EXP}: 27.1mm ’ .‘LBb‘ULTd rE-LOLI TEE Fl GEE-EhATI 01.: ........... 9 BREEDING BEHAVIOR II THE r‘a GEhERAi ION ----- . ......... 12 The 9:7 ratios .-. ...................... 12 The 27:37 ratios ---------------------- 13 The 9:7 and 27:57 ratios ------------------ 13 THE CONSTITUTION 02‘ THE b‘ThAINS USED IN THIS IlW’I-Ib'DIGcKTION - --- 18 BREEDING BEHAVIOR 1N F3 (5151553me ................ 22 Results of progenies from the 9:7 F2 group - - - - - - --- 22 Results of progenies from the 27:37 32 group - - - - - --- 26 Segregation of yellow and white endOSperm characters - --- 29 TESTIHG THE E3 FOR Si Lithins OE A_"“SlER IELLO.I DEIIT CORN - — ——— 39 Results of the seedling test ---------------- 59 Results of crosses between F A—, 0-, and R-tester types and Emerson's A-, 0-, and R—testers ---------- 45 DESCRIPTION Aim ERECTION OFS Tmms 0F A-‘i‘h "223.1 YELLO".I . :IIIT 00.111 ----------------------- 57 Ram m: 0: THE across 158‘ AND rr TO TEE DDV'HLO .11.. hm or mums mm: PI‘ “25': as rouzu: IN r5515 15:23:51.3": . --— s7 Seedling color in the Pl and El generations — - - - - - --- 67 Seedling behavior in the F2 and F3 generations - - - - --- 69 F3 seedling behavior in the 9:? Fa group .. - - - - --- 75 F3 seedling behavior in the 27:37 E3 group - - - - --- 87 MY - - - - - - «- ---------------------- 99 BIBLIOGRAPHY --------------------------- 103 - -. .- -7 ... .... _ ,. ._. r .- ‘, —>~ .. —- .- _ — -— — .. -- —— .— H v H -~— .- . h —— —— - -- 7— ._ H , , .. . - _—-- _ .. .. . n . — - .—. o— _. - H . . ._—-.- c. \ _.—. - - - _. . . - —-.-. ,. . .. . .— - -- _. ., — .. —— _ Q I —- .—. —. -- . . -— ~ I \ . . . a. .- — - - — ~ . - . .. -. .. rw ' . o -. a .. ~— , t . ‘ . ,.. _, . -_ -. — , - —. -- .- .- _ -1- INTRODUCTION The purpose of this work was to develOp pure strains of a new corn called "A—tester Yellow Dent” to be used in the production of a possible commercial variety of such corn for the liichigan farmer. Such strains have already been derived by this eXperiment from crosses made between sane of the strains which were used here for the corn breeding work and the A-tester corn of Professor R. A. Emerson of Cornell University. The latter is an inbred white flint corn having the genetic constitution aaCGRIer, while the former are yellow dent corns whose compositions were found to be either AACCrrYY, or LACcrrYY, or “con-Yr. Here A is the anthocyanin factor, dominant to its recessive allelanorph(a), and aids in the develoynent of color in the aleurone layer of the seed when the other factors 0 and R are present. The 0 factor dominant over its partner (c) is said to be a chromogen, a basic substance which, when acted on by the enzyme B in presence of A, is oxidized to a red color and to a purple color if another dominant factor, Pr, is present. The seeds, whose aleurone layer is either red or purple, will be termed “colored" in this thesis. If either A, C, or R is lacking, the aleurone layer is colorless, and hence these seeds will be designated as "colorless". I and y are a factor pair which controls the yellow and white endosperm color, the former being dominant. These strains of A-tester Yellow Dent corn have the composition -2- aaCCRR inherited from the A—tester and the yellow dent character (YY) from the mother strains. By having such censtitution (aaCGRRYY), they can not escape from observation whenever they are contaminated with any common corn which contains the A factor. If they are pollinated accidentally, by pollen from any common corn in a neighboring field, the hybrid seeds will become colored due to the presence of A, C, and R factors where A comes from the pollen parent whether it has yellow or white endosperm. The hybrid seeds can be eliminated simply by discarding the colored seeds. The colorless ones can then be saved since they are always pure. As it is now, if a pure yellow variety is contaminated by being pollinated with a white type, the grower can not know the same year that this has actually happened in his corn, as all hybrid seeds are yellow. He will not realize this situation until the next year's crop develOps, when he sees a number of white grains appearing in his yellow corn, due to natural chance recombination. Under these circumstances, he will have a very hard time to eradicate all the white seeds because the yellow endosperm character is dominant over or covers up the white one and thus the white seeds will reappear each year whenever the yy condition is effected. His pure variety of yellow corn will deteriorate as seed stock. Moreover, a contamination like this is a menace to his business, if the farmer is a seed-corn grower, since it will make him lose the confidence of his customers and hence, his market. -3- Also the contaminated ears, in most cases, could not get the first place or grade in the market and in shows. From this point of view, these newly develOped strains, if they produce a promising variety, may benefit the farmer a great deal and solve this difficult problem of contamination. HATERIALS.AND METHODS USED The work was begun as a problem of the Emperiment Station here by the late Professor F..A. Spragg in the year 1925 and was carried on by his associates in 1924 and until all materials were transferred to the writer in the fall of 1925. it is with gratitude that the writer acknowledges his indebtedness to Professor Spragg who was the originator and who laid down the foundation for this work. Strains used in this investigation as the maternal material are listed in Table I-a. Three strains of M..A. C. (Ac 254), one strain of Michigan White Cap K Bailey (Ac 256), four strains of Duncan (Ac 234), and five strains of Bailey (Ac 128) are yellow dents with red cobs. There are two strains of M. A. C. and one strain of Nelson K Bailey (Ac 255) which are dented, red cobbed, but with red pericarps. One strain of Duncan (Ac 235),and seven strains of M.A.C. (Ac 254) which came from a cross of Jhite Cobbed Golden Glow x white Cobbed Duncan, are yellow dented but white cobbed. All these strains were inbred or selfed for one or more years before used for this work as is shown in Table 1-8. The L—tester corn, which served as the paternal material, was obtained from Professor Enerson in 1922. It is shown in Table I-b. It is an inbred white flint corn with white cob and has been used by Professor Enerson and his students for testing the aleurone color factors. Its genetic constitution is definitely known as 13.8.00115111.1 At the same time, the so-called G-tester and B-tester were also received. These testers have been named for the recessive factor present; consequently, the C-tester would be uccRR, and the R-tester, uOGrr. Both are white dented and white cobbed. . The plant color of this Latester and of this c-tester was green throughout the plant body. In other words, there was not even a trace of anthocyanic pignent in any port of these plants. It was proved that the A-tester carried the allelomorphic factor R838 as well es as which causes no pigmentation (pages 67-99 ). The G-tester was found also to carry £833 and hence it was non-pignented. Some of the B-tester plants showed some color in stalks, silks, glumes, and anthers, and so they carried rrrr which causes such pigment. A few R—tester plants were found to be 1.31.3. However, only the 33 factor of the A—tester is concerned in our work later on. All self- or cross-pollination was carefully controlled. Begs of lZ-pound size, square bottom, 40-pound weight No. l Kraft 1. The La, 0-, and B—tester had white endospems and hence were yy in constitution. Hereafter this endospenn factor will be disregarded unless especially mentioned. -5... TABLE I-a. Showing the names, accession numbers, selection numbers, plat numbers, kernel color, cob color, years selfed and years intercrossed of the strains used in this investigation. Source or ‘Loooss- 1922 1923 Color of Years Years Name of variety ion seleot- plat selfed inter- number ion No. No. kernel cob crossed ................. zafimu------_-_-----___- White Gobbed Duncan 235 02600 30100 yellow white 1920 .;-— White Cobbed n. A. C. . White Cobbed Duncan 235 92600 ” a White Cobbed G. Glow 239254 12901 30200 1921 1922 White Cobbed M.‘A. C. 0'254 02600 30300 " " ” " 12904 “'32- ' u ' ll .. n 1 .11 25202 30400 " " " n n a n n " 25203 30500 n 6 N n N I! OI N 0. 25503 30600 I! I. I! I. i. N I. I! I. 2550 6 30 700 H M I. N N I! I! H N 25312 30 800 N N I! '9 Red Cobbed w ”on. r: " 25205 30900 " red " " " ” " " ' 25206 31000 red "' " " n " " " " 25207 31100 yellow N n " I' N I! N N 25502 31200 IO N N I. " " " " " 25313 31300 red " " " Nelson K Bailey 128255 23000 31400 to n ;2;7-;9L2 1921 233 1920 Michi an White ca d g x.231££; 122256 23100 31500 yellow » 1211:1212. n 137 1917-1920 Red Cobbed Duncan 234 23400 31600 “ " 1920-1922 " " " " 23701 31700 " " " I! N fl 0. 25705 51800 N '0 fl " " " " 23502 31900 " " " Source or Name of variety TABLE I-a continued. -6- .1922 . 1923 Color of .Years ,Years selfed inter- akernel [cob crossed TABLE I‘bo kernel color Name of variety .Access- ion select— plat ,Number ion No. No. or Plat NO. 128 24101 32000 24202 32100 24204 32200 24301 52300 24302 52400 yellow red 1917-1919 - 1922 " " 1917-1920 1921 1922 " “ 1917-1919 1922 " “ 1917-1922 " " 1917-1922 Showing names, accession numbers, plat numbers, and cob color of Emerson's aleurone-color testers. Accession Number 1923 Plat NO. Color of Kernel Cob A-tester C-tester R-tester 247 248 249 52500 32600 32700 white white I. N -7— paper were used for covering the tassels while transparent glassine paper bags (6" x 2%”) were used to cover the ear before the silks emerged. Two men, working together, did the pollinating at the time of crossing and selfing. One man took off the tassel bag with its fresh supply of pollen grains and shook them onto the silks cautiously, as soon as the other man quickly tore off the top of the ear bag. The latter fastened the tassel bag over the ear with No. 3 gem paper clip, while the former labeled the plant, crossed or selfed, with a paper tag. Only one pollination to an ear was made, in order to diminish the chances of contamination. Notes were taken on plant characters and growth in the field. After harvest counts of colored and colorless losrnels on the eelfed ears were made in the laboratory and then these ears were classified into groups, according to their Mendelian ratios. Notes were taken on ear characters, such as: color of kernels, color of cobs, length of ear, length of kernels, indentation, circumference of tip and butt, weight of ear, and number of rows, etc. All these notes were entered into the selection book together with the preper selection lumber of the ear. It might be well to say here a few words concerning the recording system which is used by this station, in order to help the reader trace the records in this paper. The system consists of three parts, i.e., the accession book, the plant breeding register, and the selection book. Each crap has its series of accession numbers. Any material -8- of one variety or one cross collected from one source has an accession number'aseigned to it according to the order in which it is obtained. In the accession book, besides the accession number, the name of the variety, from whom received, source, date received, and date of enter- ing the nursery, are also recorded for each.material. Before planting time, each lot of seeds from.one variety or from one selected ear is registered in the plant breeding register and given a plat number. The plat number consists of five or more figures. The first or left hand figure indicates the year in which the material is harvested. The last, or right hand, two figures designate the number of individual selections, up to 99 which are made in that plat. The digits between the first one and the last two figures indicate the number of that particular plat. Ebr instance, 26306 in Table I-a is the selection number of the sixth selection of plat 53 of the crap year of 1922. If the number of selections exceeds 99, other plat numbers are assigned to that plat to care for the extra selections. In the breeding register, the plat number, the selection number, if any, and the accession number, are all recorded in order to identify the individual material in the test. In the selection book only the individual selection number and accession number are noted. This system has been followed throughout this experiment. .A word of explanation is necessary regarding the methods used in calculating the biometrical constants given in this paper. -9- The formula used for probable error of a Mendelian ratio is P. E. = 0.6745 pqn , where n is the total number of individuals in the eXperiment, p is the percent of one member of the ratio, and q is the percent of the other member of that ratio. Thus in a 3:1 ratio, p g 75% (.75) and q z 25% (.25); and in a 9:7 ratio, p _-_-, 55.25% (.5625) and q = 43.75% (.4375) and so forth. The probable error detemined by this formula was compared with the actual deviation from the theoretical eXpectancy. For tests of closeness of fit in cases where the number of classes was more than two, use was made of the method known as X2 suggested by Harris (1912). For some statistical study in this work the following formulas were used: ti 3 zgv Here, M 8 mean; V 3 a variable; f a class n frequency; 5. = summation; n = total number of variables. qr"; (id: 0' = standard deviation; d a deviation n from the mean. 0,7,; 0" x 100 0.7. g coefficient of variability. ll E,3 _-.- 0.67450“ Es : probable error of single determination. Em -_-_ .QJIM Em 2 probable error of mean. n E; - 3 3 BA 1 3:; EA - 5‘ = probable error of difference be- tween variables A and B if no correlation exists between these two variables. -10... smmmm RE'oULTS 1311012 THE F1 GL‘I‘MIATION In the summer of 1923, twenty-four strains of different varieties used as the female parents (P1) were crossed in the field with the A-tester, which served as the pollen parent. Six crosses were made with each strain. The results are listed in Table II. In the same way these strains were crossed with the C-tester and with the R-tester. Unfortun- ately, the exact results from these two series of crosses were lost. As indicated in Table II, all these strains X the A-tester, except one strain, plat 31500, which failed in cross-pollination, resulted in a total of 64 hybrid (F1) ears all of whose kernels were colored. The color of these F1 ears was caused by the complementary action of the three dominant aleurone color factors A, C, and R. The factor A in the hybrids must have come from the side of the female parents since the male parent, A-tester, is known to have contributed no A factor but ongthe C and R factors to its off- spring. Therefore, all the female parental strains, used in the test, contained AA in their constitution. Information1 obtained indicated that all hybrid ears result- ing from crosses between the strains in plats 31400 and 31700 to 32400 with the C-tester (AAccRR) were colorless. Obviously, these strains must have been co in order to produce the colorless pro- genies, because A and 11 without C will not give the colored aleurone 1. Such information was furnished by Mr. H. M. Brown, Research- Assistant in the harm Crops Department of the Michigan State College. k...“- ‘.-. .... e . TABLE II. -11— Showing the number of colored hybrid ears (F1) which resulted from crosses of the strains used in this experiment with the A-tester, and their F1 selection numbers. Source or Name of Variety Access- ion No. 1923 Plat NOe or Female parent A- NOe Of tester colored or male produced parent ears by 01‘08868 Selection No. of £3 ears White Cobbed Duncan White CObbGd EJeCe n n n ..H H ' n n n n n n n a ne u n n n n n u n u u n n n n n n n n u Nelson K Bailey Mich. Wh. Cap K Bailey Bed Cobbed.Bunesn I! II It ‘0 It It It It It Bailey I! I! I. I! Total ............... N 256 128 (A O ,p O 0 >1 30500 30700 30800 30900 31000 31100 31200 31300 31400 31500 31600 V1 >< ti V: #4 >4 >4 >< >4 >4 P< >4 k< 31700 31800 M 31900 32000 32100 32200 32300 X 32400 X >4 #1 >4 >4 I. .. It I. N at N IONUONNUOU Pub 33 301150,-151 30235,—56 30331,-32,-33 30408,-09 30513,-le,-15 30604 30702,-03 30806,-07 30904 31008,-09,-10 31111,-12,-13 31203,-04,-05,-06,-07 31306,-07,-08,—09 31430,—31,-32 31604,-05 31703,-04 31802,-03 31901,—02 32001,-02 32101,—02 32201,-02 32301,-02 32407,-03,-09,-10 -11£L-‘ in the hybrid seeds. 0f the hybrid ears resulting from crosses of the strains in plats 30100 to 31300, 31500, and 31600 with the C- tester, some were all colored and some were only partially colored. This indicated that some of these strains were 00 and the other Co. The information also stated that these 24 strains X.theLR- tester (AACCrr) resulted in only colorless hybrid cars. Evidently, all these strains lacked the factor B and hence they were rr. The aleurone color can not be produced in this case as only two dominant factors, A.and C, were present in the hybrids. Summing up the above: all these strains used in this investi- gation are supposed to be of three types - some of them.AACCrr, some AACcrr, and some AAcorr. The whole situation can be understood more clearly by the following factorial diagram, which is in accord- ance with the above observations and verified by the results of the Eh and F3 generations. Types of these strains used in this investigation. Paternal parent maternal parent AACCrr AACcrr AAccrr Progeny tA-tester - aaCCRB Colored Colored Colored C-tester - AAccRR Colored Half colored Colorless Half colorless R-tester - AACCrr Colorless ’ Colorless Colorless -12... BREEDING BEHAVIOR IN THE F2 GEREMTION Fifty-four F1 ears were selected. Seeds from these ears were planted in 54 plate and self-pollinated in the summer of 1924. It was found that some of these plats gave ears only with 9:7 ratios or 27:37 ratios of colored to colorless seeds while the other plats gave some ears with 9:7 and other ears with 27:37 ratios of colored to colorless seeds. They are dealt with.separately in the follow- ing sections. The 9:7 ratios Twenty-nine selfed ears from.3 plate were found to have had 9:7 ratios of colored to colorless kernels. The results are given in Table III-a. A total count of segregates on these ears is as follows: -Colored- -Colorless- -Total- Observed 6780 5220 12000 Expected 6750 5250 12000 Deviation 4' 30 - 3O 0 .2!Ia_. : {3929. e 0.8 P. E. 35.5 The deviation is only 0.8 times its probable error. Therefore, the observations are very close to expectancy. The fact can be explained on the theory of complementary factors. It has been pointed out that aleurone color deveIOps only in the presence of all three dominant factors.A, C and R. .A 9:7 I: -13- ratio should result when the Fi is heterozygous for two factors. One recessive factor (a) naturally comes from.the male parent, Aptester (aaCCRR), and the other (r) from the female parent. The constitution of the mother strains (P1) must have been.LACCrr in this case, whence the 31 genotype was AaCGRr. Theoretically, when such an ear is self- pollinated, on the Eh ear 9 kernels, having all the dominant factors, would be colored, and 7 kernels would be colorless, since only one or two or no dominant factors were present. The 27:37 ratios Two hundred and five F2 ears from.28 plats exhibited a 27:37 ratio of colored to colorless seeds. The results are summarized in Table III-b. The total number of seeds on these ears is given below: -Golored- -Colorless- -Total- Observed 33823 46510 80333 Expected 33890.5 46442.5 80333 Deviation ~67.5 '*67.5 0 ._2§14_.: élafi : 0-7 P. E. 94.4 The deviation is only 0.7 times its probable error. Obviously, observations accord with expectations very well. Explained on the theory of three complementary factors, the F1 genotype should be heterozygous for the three aleurone factors, i.e., AaCcRr. Here the recessive factor (a) obviously comes from the Aptester (aaCCRR), and (c) and (r) from the mother strains (P1) which must be AAccrr in this { -14- TABLE III-a. Showing the lots which produced only F2 ears with 9:7 ratios of colored to colorless kernels. These ears resulted from the F1 genotype AaCChr (AACCrr X.aaCGRh). Name of variety .Access- F1 Po Number of ion 1923 1924 927 No. Selection Plat ears NO. NO. ‘ H.l.0. Yellow Dent 254 30331 41100 20 .. fit . . H. ' It - I]. 30513 41600 5 n N " " " 51203 43100 4 Total ------------------ 29 TABLE III-b. Indicating the plate which produced only F2 ears with 27:37 ratios of colored to colorless kernels. These ears resulted from the F genotype.AaCcRr (AAccrr X aaCGRR). 1 Name of Variety Locess- F1 F2 Number of ion No. 1923 1924 27:37 Selection Plat ears N0. N0. {his} 6031;331:123; ' ’ £3; " '361156 ' ' 20306 """ if " " who. Yellow Dent 254 30255 40900. .2 “n. 4 ‘vn- '.ar .0. 30703 42100 3 T " " a 31111 42800 5 T " ” “ 31113 43000 4 u " " " 31207 43500 7 ' n ” " 31306 43600 3 " " " " 31307 43700 3 " " " " 31309 43900 21 nelson X Bailey 255 31430 44000 8 " ” " 31431 44100 7 T T ” ” 31432 44200 8 ._ .- _ _. ~ ,_ - -. _. _ -. C u» .— “ a. 4a hr ’-- . .~ x ‘ .K ~ 0 - .. . v ‘ . . v. .— a. n- -‘ “- a—u ‘ ’ — r.- U— Q. I e 0 A . 1 I O 7 ' 1 0 1' Y ,1 o u I t I 1 - -' i v Name of variety Red Cobbed Duncan Total ------------------- -15.. TABLE III-b (continued). Access- ion NO 0 128 1923 Selection N00 31605 31703 31704 31802 31803 31901 31902 32001 32002 32101 32201 32202 32301 32302 32407 32408 1’2 1925 Plat NO. 44400 44500 44600 44700 44800 44900 45000 45100 45200 45300 45500 45600 45700 45800 45900 46000 Number of 27337 Bars 14 19 14 12 417 205 -15- case. When such an F1 genotype is self-pollinated, on an Eh ear 27 kernels, having all three dominant factors, would be colored, and 37, having at least one homozygous recessive pair of aleurone factors, would be colorless. . The 9:7 and 27:37 ratios Twenty-one plats, each from a single F1 ear, prouuced ears with 9:7 ratios and ears with 27:37 ratios of colored to colorless seeds. Out of 200 such F2 ears, lOl.were found to have 9:7 ratios, 99 to have 27:37 ratios. The results are summarized in Table III-c. Counts for these two sorts of ears gave the following numbers of kernels: -From 101 9:7 F2 ears- -From 99 27:37 32 ears- -Colored- -Colorless- -Total- -Colored- -Color1ess- —Total- Observed 20834. 15958. 36792 18215. 24588. 42802 Ripected 20695.5 16096.5 35792 18057.5 24745.5 42802 Deviation +1ss.5 -l38.5 0 41.57,! 457.5 0 211:. = m a 202 151.! = 203 Page 64.2 68.95 The deviation for the first lot is 2.2 times its probable error and thus indicates a 9:7 Mendelian ratio. The deviation for the second lot is only 2.3 times it probable error, obviously indicating a 27:37 ratio. Glancing at Table III-c, it is interesting to note that 9:7 ears and 27:37 ears, which occurred in each plat, were in equal numbers. ‘A calculation on this basis for the total number of ears in the 21 plate shows a deviation of 1, which is only 0.2 times its probable \J. - A. e, Name of variety Wh. Oobbed Duncan 1!. A. 0. 0'. H I. “-313 111-0 0 and 27:37 F3 031... -17- Showing the plate which produced both 9:7 They resulted from the F1 genotypes AaCCRr and AaCcRr which occurred on the same ear (AACcrr X.aaCCRR). Red Cob. Duncan 234 Acce5&- Fl 32 Actu81.No. Theoretical ion 110. 1923 1923 of ears in No. of ears P21- Selec- plat (1:1) in Dev. P.E. P.E. tion No. 9:? 27:37 '9:7 27:37 No. ___________________ ---------- 235 - -301151 -40800- - 4 3 3.5 3.5 0.5 0.9 0.6 254 30256 41000 4 9 6.5 6.5 2.5 1.2 2.1 T 30332 41200 12 4 8.0 8.0 4.0 1.4 2.9 T 30333 41300 3 11 7.0 7.0 4.0 1.3 3.1 v: 30408 41400 9 7 8.0 8.0 1.0 1.4 0.7 T 30409 41500 4 7 5.5 5.5 1.5 1.1 1.4 T 30514 41700 5 6 5.5 5.5 0.5 1.1 0.5 T 30515 41800 7 9 8.0 8.0 1.0 1.4 0.7 T 30604 41900 1 4 2.5 2.5 1.5 0.7 2.0 T 30702 42000 2 l 1.5 1.5 0.5 0.6 0.9 T 30806 42200 4 3 3.5 3.5 0.5 0.9 0.5 T 30904 42400 1 5 3.0 3.0 2.0 0.8 2.5 T 31008 42500 8 13 10.5 10.5 2.5 1.5 1.7 T 31009 42600 4 4 4.0 4.0 0.0 0.9 0.0 T 31011 42700 7 7 7.0 7.0 0.0 1.2 0.0 T 31112 42900 4 1 2.5 2.5 1.5 0.7 2.0 T 31204 43200 3 2 2.5 2.5 0.5 0.7 0.7 T 31205 43300 2 4 3.0 3.0 1.0 0.8 0.1 " 31206 43400 5 1 3.0 3.0 2.0 0.8 2.5 ” 81308 43800 5 2 3.5 3.5 1.5 0.9 1.5 31604 44300 8 6 7.97_ 7.0 0 3 7 Total . ----------- 101 99 100.0 100.0 1.0 4.7 0.2 f! -13- error. This leads to the conclusion that the El genotypes from which these F2 ears resulted must have been of two sorts, AaCCRr, and AaCcRr, but all look alike, i.e. all are colored. Therefore, their parental (P1 or mother strain) constitution must have been “Corr. When crossed with the L-tester (aaCCRR), two types of gametes, ACr and Acr in equal numbers from this female parent were fertilzed by the gametes, a011, _ from the L-tester. When self-pollinated, the genotype AaCGRr produced 9:7 F2 ears, while the genotype AaCcRr produced 27:37 F2 ears, in proportion of 1:1. Summing up the above: the hypothesis fits all observations in this generation very well, and also verifies the results of the F1 generation. It is further verified by the F3 generation. THE CONSTITUTION OF THE STRAINS USED IN THIS INVESTIGATION hereon (1918) has pointed out that all corn varieties which he tested were rr. In other words, they lacked the R factor. Like- wise all varieties were found to be 44 except two, one grown locally in Southern Missouri and the other in Ohio. There is apparently greater diversity with respect to the 0 factor. Some of the varieties are apparently 00, thus being B—testers, while some are Go and others cc, and yet others a mixture of these types. It has been found that all strains which have been tested in this experimental work were AA and rr but were inconsistent with regard to the 0 factor, conclusions which coincide with mnerson's findings. The results are arranged in Table IV, which is in pedigree form. By it the female parents and their progenies in the F1 constitution of these maternal Pl strains was determined by the breed— and F2 generations can be traced in either direction. The genetic ing behaviors of progenies of the 31 and F2 generations. Glancing at Table IV, one will see that strains of White Cobbed Duncan (Ac 235) and Red Cobbed Duncan (Ac 234) had the constitution of.AACcrr or AAccrr. Strains of'M. A. 0. (10 254) had AACCrr, AACcrr or AAccrr. This diversity of the Co factor pair may be due to the fact that some of the strains had been either crossed or intercrossed and self- pollinated previous to this investigation. For such information the reader is referred to Table I-a. The strains wdth Co in composition would keep on segregating in a simple Mendelian fashion (3021c), whenever they were self-fertilized. Probably this is what has actually happened in nature, and has kept nearly all common corn varieties so diverse with regard to the C factor. Fer some unknown reason such heterozygous condition has been rarely found for the A and r factors. Bailey'(Ac 128) and Nelson K Bailey (Ac 255) have the constitution AAccrr. ""EDIIIG BEIQVIOR III THE ‘5 GBI-IERATION For the 1925 (33) planting eighty-one F2 ears were selected; of these 29 had 9:7 ratios and 52 had 27:37 ratios. Only colored TLBEE IV. flame of Variety Th. Cob. Duncan Who CObe Duncan 'l'lhe 00b 0 Ge Glow M.A.C. 1 Access- ion No. -20.. P 19%. selec- tion NO 0 25202 25203 25303 25306 25312 25205 25206 25207 F1 1923 selec- tion No. 31311.51 30255 30256 30331 30332 30333 30408 30409 30513 30514 30515 30604 30702 30703 30806 30904 31008 31009 31011 31111 31112 F 195; plat \v 1.0. 40700 40800 40900 41000 41100 41200 41300 41400 41500 41500 41700 41800 41900 42000 42100 42200 42400 Genetic constitution of the maternal P strains used in this investigation. Genetic constitution AAccrr ARCcrr AACCrr AACcrr a .3 AAccrr AACcrr AACcrr AAccrr AACCrr era *-21- sTABLE IV. (continued) 32408 46000 ‘Name of Variety (Access- Tl F1 'F2 ‘Genetic ion No. 1922 1923 1924 constitution selection selection Plat NO. NO. NO. #4- .Fi'.. 4.. 2.54 31113 43000 Mccrr (continued) 25302 31203 43100 AACCrr 31204 43200 AACcrr 31205 43300 T 31206 43400 a 31207 43500 AAccrr 25313 31306 43600 T 31307 43700 T 31308 43800 AACcrr 31309 43900 T Nelson K Bailey 255 23000 31430 44000 leorr 31431 44100 T 31432 44200 T 223400 31604 44300 AACcrr _ 31805 44400 " Red Cob. Duncan 234 ¥}23701 31703 44500 AAccrr ) ; 31704 44600 T f '23705 31802 44700 T 7 31803 44800 " ‘23802 31901 44900 T 31902 45000 T Bailey 128 24101 32001 45100 " 32002 45200 T 24202 32101 45300 T 24204 32201 45500 T 32202 45600 T 24301 32301 45700 T 32302 45800 T 24302 32407 45900 T -22.. kernels from the two groups were planted and selfed in the field, while the colorless ones from such ears were discarded. The reason for planting the colored individuals and discarding the colorless ones is that the strains of the Aptester yellow dent corn can.be derived more easily from the former than from the latter in terms of possibility and economy of work. Iheoretically only 1 out of 7 or 37 colorless kernels on a 9:7 or 27:37 ear, reapectively, is the A—tester type, while out of 9 or 27 colored kernels from the ’37 or 27:37 ear, 4 or 6 will, when self-pollinated, produce 3:1 ears of which one-half or one-third of the colorless kernels will be the.A-tester types looked for. (See factorial diagrams on pages 23 and 26.) Therefore, using the colored individuals of F2 ears will save the plant breeder more time and will give him more chance or possibility of develOping strains of the A-tester yellow dent corn than if he uses the colorless ones. Segregations for aleurone color of all individual F3 ears were counted. All these ears have been grouped into two groups, i.e. those resulting from the 9;? F2 ears and those resulting from the 27:37 F2 ears. Each group has shown similar consistent results. Results of pr0genies from the 9:7 F2 group Three kinds of mendelian ratios for F3 ears have been found in any plat of this group. In each plat, some ears had 1:0, some had 3:1, and others had 9:7 ratios of colored to colorless seeds. A summary of data for this group is set forth in Table v;s. Theoretically, colored seeds of any 9:7 F2 ear, if from a cross of uCGrr X aaCGRR, should act in the following manner when they are selfed in the F3 generation. _ -Phenotypic -Fé genotypes— -Breeding behavior in F3- PrOportion of F3 ears- 1 AACGRR 1:0 (only colored) breeds true 1 2 AaCCRR 3 colored: l colorless (aaCChh) (A-tester type) 4 ZMCCRr 3 colored: l colorless (AACCrr) (R-tester type) 4 AaCCRr 9 colored: 7 colorless 4 Such was proved to be the case by the actual observations. As seen from Table V-a, the numbers of ears in each plat with these three different Mendelian ratios were in the pr0portion of 1:4:4. Values of X2 for most plats were less than 1, hence the value for closeness of fit (P) is very high. Evidently observations are very close to eXpectation. The total numbers of actual and expected ears of these classes from 29 plats are given below: Ratio 1:0 3:1 9:? Observed total 53.0 233.0 230.0 Expected total 57.3 229.3 229.3 ' Deviation -4.3 + 3.7 v 0.7 X2 for. total = 0.3880 P = very close fit. There were two kinds of 3:1 ears: one resulted from the F2 genotype “COBB, and the other resulted from the F3 genotype MCGRr. The first kind of 3:1 ears is the one which has been looked for by this experiment, because its colorless seeds are the A-tester type F2 1924 Selection N00 TABLE V;e. F3 1925 Plat N00 -24’ number of 33 ears, resulting from.colored in- dividuals of 9:7 Fa ears, showing 1:0, 3:1 and 9:7 ratios. Actual No. of ears 40807 41102 41109 41113 41114 41118 41119 41201 41211 41212 41216 41311 41409 41413 41503 41512 41602 41604 41605 54000 54100 54200 54300 54400 54500 54600 54700 54900 55000 55400 55500 55600 55900 56000 56100 56200 180 3:1 9:7 0 3 6 2 8 9 3 17 12 3 7 8 2 18 13 2 5 7 3 10 7 2 6 7 2 13 _6 3 11 10 2 7 9 4 11 9 3 7 11 3 10 8 1 13 10 1 11 9 1 11 10 3 7 6 1 6 6 1 7 6 Theoretical No.(1:4:4) of cars with ratios of with patios of 1:0 3:1 9:7 X2 Closeness of fit (P) 1.0 2.1 3.6 2.0 3.7 1.6 2.2 1.7 2.3 2.7 2.0 2.7 2.3 2.3 2.7 2.3 2.4 1.8 1.4 1.6 4.0 4.0 8.4 8.4 14.2 14.2 8.0 8.0 14.7 14.7 6.2 6.2 8.8 8.8 6.7 6.7 9.3 9.3 10.7 10.7 8.0 8.0 10.7 10.7 9.3 9.3 9.3 9.3 10.7 10.7 9.3 9.8 9.8 7.1 7.1 5.8 5.8 6.2 6.2 2.2500 0.0658 0.9766 0.6250 2.4620 0.4642 0.8125 0.1502 3.1070 0.0937 0.2500 0.9375 1.0714 0.4286 1.5938 1.0712 0.8739 1.0156 0.1453 0.3752 0.3318 very good 00 I. 0.3011 Very good 90 I. 0.2137 very g00d N l. I! I! 0.5895- Very good 0.4648 0.5896 Very good 0.6028 Very good -25.. TABLE V;a (continued) F 'F3 .Actual Theoretical 1922 1925 no. of cars no.(1:4:4) of ears X? Closeness Selection Plat wiih raiigs of ' with ratios of - of fit NO. No. 130 3:1 9:7 1:0 3:1 9:7 (P) 41701 56300 1 6 10 1.9 7.5 7.6 1.5293 0.4802 41708 56500 1 6 6 1.4 5.8 5.8 0.1453 vary good 41802 56600 3 6 9 2.0 8.0 8.0 1.1250 0.5767 41810 56700 1 3 10 1.6 6.2 6.2 4.1605 0.1268 41904 57000 2 6 5 1.4 5.8 5.8 0.3227 Vbry good 42203 57300 1 6 4 1.2 4.9 4.9 0.4544 ” " 42208 57400 1 4 5 1.1 4.4 4.4 0.1249 " " 42505 57700 2 4 7 1.4 5.8 5.8 1.0192 0.6020 42708 58100 2 5 9 1.8 7.1 7.1 1.1561 0.5693 29 plate Total ----- 53 233 230 57.3229.3229.3 0.3880 Very good which has the constitution of aaCCER and from which the strains of the ‘L~tester yellow dent will be derived. Colorless seeds of the other kind of 3:1 ears are the R-tester type (AACCrr). Since these two kinds of 3:1 ears all look alike, they must be separated from each other by a method which will be dealt with later on. Results of progenies from the 27:37 52 group Plants grown from colored seeds of any 27:37 F2 ear bred differently in F5, as was expected. In nearly every plat were found F3 ears with 1:0, 3:1, 9:7, and 27:37 ratios of colored to colorless seeds, but 110 ears did not appear in some plats, since they occur theoretically only once in 27 cases. The results are listed in mable VLb. 0n the theory of complementary factors, the 27 F2 colored in— dividuals, when selfed, will produce the fbllowing types and in the following preportion in F5. -F2 genotypes- -Breeding behavior in F3- ~Pr0portion of F3 .phenotypes- 1410038 1:0 (only colored) breeds true 1 2AaCGRR 3 colored: l colorless (aaCGhR) (Arrester type) ZLACcRB 3 colored: l colorless (AAccRR) 6 (C-tester type) ZLLGGRr 3 colored: l colorless (AACCrr) (Rptester type) 4LaGGBr 9 colored:7 colorless 4LaCcRB 9 colored:7 colorless 12 4LchRr 9 colored:7colorlese 84800Rr 27 coloredz37 colorless 8 F 1954 Selec- tion EABLE 74b. -37- Number of F3 ears, resulting from colored in- dividuals of 27:37 E: ears, showing 1:0, 3:1, 9:7 and 27:37 ratios. F5 1925 Plat N00 Actual No. of page wiin gagigs of 1:0 3:1 9:7 27:37 Theoretical No. (1:6:12:8) of ears with ratios of 1:0 3:1 9:7 2773? Close- ness of fit 44505 44511 44505 44509 44514 44711 44507 44905 40701 40702 40705 40705 40705 40709 40710 40502 40505 40901 51700 51800 51900 52000 52100 52200 52300 52400 52500 52600 52700 52800 52900 53000 53100 53200 53300 53500 0 5 8 0 5 9 1 7 9 2 3 10 1 8 10 1 3 9 3 4 12 0 5 10 0 s 7 0 8 13 0 6 11 0 3 3 1 8 14 1 2 14 0 3 7 2 1 5 0 3 11 0 7 8 0 2 9 1 6 6 1 5 10 0 2 6 7 0.7 5 0.7 6 0.9 7 0.8 5 0.9 7 0.7 7 1.0 10 0.9 5 0.6 9 1.1 6 0.9 3 0.3 9 1.2 10 1.0 5 0.6 5 0.5 6 0.7 6 0.8 4 0.6 4 0.6 6 0.9 5 0.5 4.4 5.8 5.6 0.3 6.7 5.1 2.0 7.1 3.3 2.9 4.4 4.7 3.3 3.8 5.3 2.9 8.4 10.2 9.8 10.7 8.9 11.6 11.1 6.7 13.4 10.2 4.0 14.2 12.0 6.7 5.8 8.9 9.3 6.7 7.6 10.7 5.8 5.6 6.8 6.5 7.1 5.9 7.7 7.4 8.9 6.8 2.7 9.5 8.0 4.4 3.9 5.9 6.2 4.4 5.0 7.1 3.9 0.9512 0.9595 2.5145 2.0195 0.7552 4.9579 2.0001 0.5751 1.5575 1.1551 1.1255 0.1550 5.4999 0.5751 5.4720 1.7125 2.2715 1.9501 2.4705 0.2547 1.1057 0.6654 0.5689 Very good 0.1731 0.5724 Very good 0.7126 0.7639 0.7730 Very good 0.3262 Very good 0.0932 0.6382 0.5233 0.5839 0.4917 Very good 0.7771 F ’ 1954 Selec- tion Io. F5 1925 TABLE V-b (continued) 7Actua1 N0. of plat ears with ratios of No. 1:0 3:1 9:7 27:37 40902 41001 41002 41213 41312 41313 41401 41504 41507 41706 41816 41902 41905 42002 42401 42501 42516 42602 42701 42802 42906 53600 53700 53800 54800 55100 55200 55300 55700 55800 56400 56800 56900 57100 57200 57500 57600 57800 57900 58000 58200 58300 1 4 11 5 1 5 10 6 Theoretical No. (1:6:12:8) 0f 2 Closeness gaze with 255105 of {X of 1:0 3:1 9:7 27:37 Fit (P) 0.5 3.1 6.2 4.1 2.8699 0.4152 0.4 2.4 4.9 3.3 5.2613 0.1562 0.6 3.8 7.6 5.0 0.9766 Very good 0.7 4.0 8.0 5.3 1.5624 0.6724 0.7 4.0 8.0 5.3 0.5625 very good 0.8 4.7 9.3 6.2 0.6964 " " 0.7 4.4 8.9 5.9 0.8679 " " 0.8 4.9 9.8 6.5 0.1138 " " 0.7 4.2 8.4 5.6 0.5396 " " 0.4 2.2 ‘4.4 5.0 1.5129 0.5154 0.5 3.3 6.7 4.4 0.7498 Very good 0.7 4.2 8.4 5.6 0.2435' " " 0.6 3.6 7.1 4.7 3.7151 0.2986 0.6 3.1 6.2 4.1 1.0268 0.7952 0.6 3.3 6.7 4.4 3.2248 0.3624 0.7 4.0 8.0 5.3 0.5000 Very good 0.7 4.0 8.0 5.3 0.5625 ' " 0.4 2.2 4.4 2.9 5.1883 0.1605 0.6 3.5 6.7 4.4 0.9749. Very good 0.7 4.2 8.4 5.6 0.7764 Very good 0.7 4.0 8.0 5.3 2.5623 0.4708 F5 1925 plat R00 EABLE 74b (continued) Actual N0. of -39.. Iheoretical N0. (1:6:1238) of ~eags with gatios of oars with ratios o; 1:3 3:1 9:7 27:37 1:0 3:1 9:7 27:37 43603 43803 43916 44001 44004 44101 44104 44207 Total 58400 58500 58600 58700 58800 58900 59000 59100 59200 52 plat835 1 1 5 6 221 8 3 3 2 15 4 11 6 14 7 9 8 7 6 5 4 8 5 448 276 0.8 0.3 1.1 0.8 0.9 0.9 0.6 0.6 0.7 4.7 2.0 6.7 4.7 545 5.1 3.8 3.3 4.4 9.3 3.9 13.3 9.3 11.1 10.2 7.6 6.7 8.9 6.2 2.7 8.9 6.2 7.4 6.8 5.0 4.4 5.9 36.3 217.8 435.6 290.4 5.9461 2.8335 4.5769 1.1785 1.7550 0.3571 1.2348 1.6494 0.7871 1.1607 0.1150 0.4218 0.2098 0.7572 0.6286 Very good 0.7476 0.6528 Very good 0.7645 -30... As seen in flable V;b, of a total of 980 ears from 52 plats, 53 were 1:0 ratio, 221 were 3:1 ratio, 448 were 9:7 ratio, and 276 were 27:37 ratio for colored to colorless seeds. Their expected numbers on a basis of 1:6312:8 ratio were 56.5:217.8:455.6:290.4. The value of 22 is 1.1607 and hence the value of closeness of fit (P) is 0.7645. This indicates that these observations are very close to GXpectancy. Here the 3:1 ears were of three sorts genotypically. Colorless individuals (kernels) of the 3:1 ears derived from the £3,3enotype AaCGRB.were the.&~tester types which were sought for by the present experiment. Likewise, the colorless kernels from the 3:1 ears of the F2 genotype AaCcRR would produce a C—tester type, Anccflfl, and the colorless kernels of the 3:1 ears of the F2 genotype AACGRr would produce an B—tester type, AAGCrr. How to sort the A-tester type out of the other two types will be described later on. Segregation of yellow and white endOSperm characters As pointed out previously, all the mother strains had the yellow endOSperm character and were crossed with Emerson‘s Aptester which had the white endosperm character. 0n the basis of a single pair of factors, the fonner should be YY'and the latter yy regardless of the aleurone factors. All the E1 kernels should be yellow due to the factor Y being dominant over its allelomorph y, and they should segregate in the manner of 2 yellow to 1 white in Ea‘when self- pollinated. It should be recalled that the yellow endosperm was wholly covered up in the F1 and partly so in the 32 by the colored .—31.. aleurones. No data were taken for these two generations. VThree groups of ears segregating for the yellow and white endosperms were observed in the colorless aleurone individuals of the 3:1 33 classes. These groups were those which bred true for either yellow or white endosperm.and those which did not breed true for the yellow endosperm but segregated in a 3:1 ratio. The results are arranged in Table VI. Ebrty E3 ears with a total of 3370 yellow kernels were found to be in the group (a in Table VI) which bred true for the yellow endosperm character, since the progenies of these ears resulted in a total of 233 ears with only the yellow endosperm in the E; generation. Thirty-five F3 ears with.a total of 2866 white seeds were noted and found to be in the group (0 in Table VI) which bred true for the [bite endosperm.but were not carried on to the EA. Eighty-two E3 ears were found to be in the group (b in Table 71) which did not breed true for either of these characters. They gave a total or 4870 yellow seeds to 1567 white seeds, where 4827.75; 1069.25 were expected. A.deviation of 42.25 was obtained, which is 1.8 times its probable error. Evidently it is a 3:1 ratio of yellows to whites on these ears. The yellow individuals were carried on to the F; generation and resulted in 141 yellow seeded ears (1:0) and 298 yellow and white seeded ears in nearly a prOportion of 1:2. Ca1- oulating on this basis the expected nwmbers for these two classes of ears are 146.3 s 292.7. The deviation is 5.3 which is only 0.8 times its probable error. Thus, observations are very close to eXpeotancy. -32- ' TdflLE VI. Segregation of yellow and white endOSperm in the colorless aleurone individuals of the 3:1 F3 class and their be- havior in E4. (8) Group which bred true for the yellow endosperm. 1925 1926 - (F3) Number or Kernels (F ) number of eazs Selection with gndosperm Plat Yellow Yellow and White NO. -Yellow- -White- No. seeded white seeded seeded 51707 90 650300 3 52603 72 650600 1 52803 30 650900 1 53005 75 651200 a 53302 105 651600 6 53303 40 651700 1 53801 83 652300 11 53915 92 652700 11 54006 86 653000 14 54116 72 653500 5 54208 14 653900 0 54304 90 654000 15 54404 80 654300 5 54509 68 654600 0 54605 101 654800 5 54608 106 654900 5 54708 92 655000 9 54906 134 655200 11 55102 98 655900 1 55202 90 656100 8 55303 70 656500 9 ‘0 -53- TABLE 71, (a) bontinued) 1925 , (33) Number of kernels Selection with endoqugg No. -Yellow- -White- 55307 146 0 55510 75 0 55804 84 0 55903 89 0 55909 40 0 56007 110 0 56303 39 0 56403 109 0 57501 105 0 57603 76 0 57901 121 0 58303 95 0 58501 62 0 58502 123 0 58602 117 0 58603 95 0 58606 58 0 58807 70 0 58702 62 0 Total 40 ears 3370 0 1926 (F ) Pla No. 656600 657100 657700 651800 658000 658100 658300 658500 659500 659700 660000 660700 661100 661200 661300 561400 661500 661600 661800 ' Epmber of ears Yellow Yellow and seeded white seeded 16 0 3 O 7 0 3 0 4 0 7 0 4 0 1 0 3 0 2 O 3 0 5 0 9 0 15 0 6 O 3 0 4 0 2 0 7 O 233 0 White Seeded -m- TABLE 71 (continued) (b) Group segregating for the yellow and white endosperms. ’ . 11:? m; 2: (1:232:13 23:? Number of ears Selection Plat Yellow Yellow and White No. -Yellow- -White- NO. seeded ‘.2hite seeded seeded 51204 63 21 650000 2 6 0 51503 81 30 650100 1 3 0 51704 72 y 20 650200 0 1 0 52301 55 25 650400 0 3 0 52504 65 21 650500 0 3 0 52702 50 21 650700 1 4 0 52703 18 6 650800 0 1 0 52902 42 21 651000 0 1 0 53002 65 21 651100 1 2 0 53202 77 25 651300 3 5 0 53205 47 14 651400 2 4 . 0 53207 15 4 651500 failed 53304 52 22 651800 2 6 0 53404 51 19 651900 3 0 0 53405 78 22 652000 1 2 0 53502 29 12 652100 0 4 0 53601 66 25 652200 0 2 0 53908 91 28 652400 5 10 0 53910 94 34 652500 6 11 0 53911 90 28 652600 0 5 0 53917 89 28 652800 1 4 0 53918 83 22 652900 4 5 0 -35.. TABLE VI, (b) (continued) 1935 Number of kernels. 1926 t (33) with endospgzg (F4) number of ears Selectionn Plat Yellow Yellow and White No. -Yellow- -White- N0. seeded white seeded seeded 54009 62 16 653100 2 3 0 54103 42 14 653200 self-pollination failed 54107 100 27 653300 2 6 0 54112 20 7 653400 1 2 0 54118 52 15 653600 1 2 0 54205 63 20 653700 failed 54207 11 5 653800 0 1 0 54308 36 14 654100 2 4 0 54310 38 14 654200 3 1 0 54405 65 17 654400 3 4 0 54408 82 24 654500 e ' 5 0 54604 59 19 654700 3 4 ‘ 0 54905 63 23 655100 0 2 0 54908 43 17 655300 3 4 0 55004 55 20 655400 2 4 0 55005 58 20 655500 2 3 0 55007 62 20 655600 2 4 0 55008 61 17 655700 0 0 0 55010 57 21 655800 0 0 0 55103 92 31 656000 4 9 0 55203 37 13 656800 0 2 0 55204 42 14 656300 1 5 0 1925 (Pb) Selection NO. 55508 55509 55511 .55603 55703 55802 55803 55904 56202 56401 56503 56801 56802 56804 56902 56903 57003 57104 57203 -Yellow— 50 65 71 62 52 86 78 90 83 60 88 66 88 59 34 49 46 50 hard to classify 35 '36- TABLE 71, (b) 'Number of kernels with endgspggg 4White- A926 (32) Plflt N00 (continued) number of ears Yellow seeded Yellow and white seeded 18 22 28 22 14 18 28 21 26 24 18 25 13 22 24 10 11 12 14 16 656700 656800 656900 657000 657200 657300 657400 657500 657600 657900 658200 658400 658600 658700 658800 658900 659000 659100 659200 659300 659400 bl NN White seeded 0 0 _TABLE VI, (b) (continued) »-37- ,1926 (FL) plat n0 0 Rumba; of ears nYellow seeded Yellow and White white seeded seeded 1925 ‘number of kernels ”‘31 W ‘Selection so. -Yellow— AWhite- 57602 55 24 57703 68 20 57802 62 22 58004 44 15 58103 70 24 58107 64 19 58108 89 _ 26 58202 36 13 58204 36 19 58402 77 25 58403 50 15 58404 83 15 58701 39 8 58704 90 25 59205 62 24 59206 hard to classify Total for 82 ears -- 4870 1567 Ripected total -- 4827.75 1609.25 Deviation total -- +-42.25 -42.25 23.4 659600 659800 659900 660100 660200 660300 660400 660500 660600 660800 660900 661000 661700 661900 662000 662100 Total for 82 plate-- Expected total Deviation total N OINN 146.3 "' 5.3 2 0 3 0 2 0 4 0 5 0 4 0 8 0 4 0 2 0 5 0 0 0 2 O 3 4 0 4 0 1 0 .5. .9. 298 0 292.7 4'5.3 DEL-251:0.8 29.3. 6.7 .lal I. ~38- mABLE 71 (continued) (c) Group which bred true for the white endosperm but not carried on to F . 1925 number of kernels 1925 number of kernels (F3) W (F3) Jul; _e_n.9._0_8_2§m__ Selection Selection, No. -Yellow— -fihite- No. ~Yellow- -Wh1te- 52701 0 138 56204 0 66 53306 0 85 56307 0 95 53804 0 83 56402 0 118 54007 0 120 56506 O 51 54008 0 62 , 56608 0 89 54108 0 58 56703 0 91 54110 0 73 56905 0 68 54117 0 92 57402 0 103 54306 0 92 58002 0 69 54505 0 123 58201 0 102 54512 0 68 58405 0 78 54610 0 31 58406 0 56 54704 0 122 58407 0 94 54803 0 50 58503 _9_ Jig 54805 0 50 Total 35 ears 0 2866 55009 0 41 55407 0 ' as 56602 0 99 56006 0 76 56106 0 71 56203 0 50 -39- TESTING TIE F3 Mi 321511133 02‘ A-TES’JSR YELLOW DENT CORN Two methods have been followed in sorting out the A—tester type from the other two types, C-tester and Retester. One method was to detect the anthocyanic pigment of seedlings grown from the colorless seeds of 3:1 33 ears. The other method was to cross these three tester types with.Emerson'std, C, and R aleurone color testers. Results of the seedling test It was thought that the.d factor was entirely responsible for the production of anthocyanic pigment in the mature corn plant and likewise in the seedlings. Consequently, the types c-tester and B. tester could be eliminated at the seedling stage, since they had the (A factor which would cause those seedlings to be colored, whereas the Iptester type could be picked out because its seedlings would be non- colored on account of lacking the.A factor. This idea was proved to be invalid, to some extent, by the results of the other method, i.e. crossing these tester types with Emerson's aleurone testers in the field (see Tables pra and IXEb). Those results have revealed that both.the.4-tester type and the G-tester type resulted from colorless seeds of 3:1 F3 ears of the 27:37 F2 group, whose seedlings sire considered to be nonscolored by the seedling test (Table prb), and that the seedling test could not be used to rogue out the c. tester type from the A—tester type. By’a later experiment (pages 67-99 ), it has been found that ~40- the multiple allelomorphic factor rr of the R series was present in the Retester type, hence the type was AACCr’rr; rr causes the seedlings to be colored in presence of the.i factor. .Also, HERE recessive to rrrr for plant color, has been found present in the.A-tester type, hence the type was aaCCRshg and in the C-tester type making it AAccRSRg. In the presence of the factor.4, ngg‘would cause these seedlings to be non-colored. This is the reason why the C-tester type could not be rogued out by the seedling test, but it could be distinguished by crossing it with Emerson's testers. In the winter of 1925, colorless seeds from 394 331 Pb ears were planted in sand trays in the laboratory for seedling tests. 0f the 394 ears, 208 were pragenies of the 9:7 Eh ears and 186 were progenies of the 2733? F2 ears. .About three weeks after planting, the seedlings were pulled out and their color was recorded. From the 937 Pa lot, of 208 ears, 99 showed colored seedlings and 109 gave non-colored seedlings. The results are shown in Table VII-s. As pointed out previously, there were two kinds of 3:1 F3 ears, genotypically, i.e. one contained the.A-tester type and the other contained the R-tester type. Theoretically, these two types should have been in equal numbers and approximately such was the case. Thus the 109 care must have been of the.A-tester type and the 99 ears of the R-tester type. They were practically in equal preportion. The expected numbers were 1043104. This exhibits a deviation of 5, which is only 1.0 times its probable error. Observations were very close to eXpectation. That this conclusion was correct was 53400 53900 54000 54100 54200 54300 54400 54500 54600 54700 54900 55000 55400 55500 55600 55900 56000 56100 a41t TLBEB VII-s. Showing that colorless seeds of 99 3:1 33 ears of the 937 F' group gave only colored seedlings, which would be of the R-tester (AGr) type; and that colorless seeds of 109 321 F ears of the same F' group gave only non-colored seedlings w ch would be of the.£-tester (aGR) type. number of 3:1 53 ears number of 3:1 33 cars giving 1925 . _____.gi_' winsw colored non-colored (33) colored non-colored seedlings seedlings Plat seedlings seedlings (AGr) (808) No. (ACr) (add) 3 0 56200 1 4 1 2 56300 3 3 ll 6 56500 2 3 2 5 56600 4 2 8 8 56700 1 1 6 0 57000 1 3 4 6 57100 1 2 2 4 57200 1 1 7 6 57300 1 4 2 3 57400 1 2 3 3 57700 2 2 5 4 58100 2 4 3 4> 58300 __3; __;L 4 5 Actual total - 99 109 3 8 Expected 1 3 total - 104 104 4 5 Deviation- -5 45 4 3 291.. = .549 = 1.04 POE. 4.8 -42— proved by the results from the field crosses between these two tester types and Emerson's testers. .As shown in Tables VIII-a, VIII-c and lips, the colorless kernels of 3:1 F3 ears, whose seedlings were shown to be non-colored by the seedling test, preduced only the A- tester type, and those whose seedlings were shown to be colored by the same test produced only the R-tester type. From.the 27:37 F2 lot, of 186 F3 ears, 63 gave colored seedlings and 123 gave non-colored ones. The results are shown in Table VII-b. (As said before, the 3:1 ears were of three sorts in this group, namely, the.A-tester type (aaCCBgRg), the C-tester type (ALccRERS) and the R-tester type (AAOCrrrr). Obviously, these 63 ears must have been the Rptester type, while the 123 ears must have been the Anteater type plus the C-tester type. Calculation on 1:2 basis showed that the expected numbers for the Ratester type and for the Anteater type plus the C-tester type were 62:124. The deviation is 1.0, which is only 0.2 times its probable error. Observation accords with expectancy exceedingly well. This conclusion was also verified by the results from the actual field crosses between these tester types and Emerson's testers, as is shown in Tables VIII-a, Vllléb,HIII-c and prb, because both the.A-tester type and the C-tester type resulted from the colorless seeds of 3:1 F3 ears whose seedlings were shown to be non-colored by the seedling test, and the R-tester type resulted from the colorless seeds of 3:1 Eb ears whose seedlings were shown to be colored by the same test. 51700 51800 51900 52000 52100 52200 52300 52400 52500 52600 52700 52800 52900 53000 53100 943- 411813 VII-b. Colorless seeds of 63 331 F3 ears of the 27237 F group produced only colored seedlings which would be of t e R-tester type (ACr); and those of 23 3:1 23 ears of the same F2 group produced only non-colored seedlings, which would be of the L-tester type plus the C-tester type. Number of 3:1 33 ears ’Number of 3:1 33 ears iving [1925 _______giving colored non-colored (F3) colored non-colored seedlings seedlings Plat seedlings seedlings (ACr) (aCR + Acl‘i) No. (ACr) (aCR «t LcR) 1 4 53200 2 4 3 2 53300 1 4 3 4 53500 1 1 1 2 53600 0 1 2 5 53700 0 o 1 2 53800 0 3 3 0 54800 1 3 l 2 55100 2 1 0 1 55200 1 3 l 4 55300 1 5 2 1 55700 1 3 1 1 55800 1 3 6 2 56400 0 3 0 2 56800 1 3 0 3 56900 1 s 0 1 57100 1 2 2 1 57500 O l 2 4 57600 0 2 1 0 57800 1 1 4 0 1 _.. s A o . - A. 1 .5 ‘ e - , . _~ . ‘ . - 1 , I . . e . ' _ . — . ' ‘ ’ g . . _ . . P r. - o 1 I . . “' ~ . , . ~ ~ ' ‘ ‘ e . I .m . . .- . ‘ , .. .. \ V v . .. , 7 . a . u .. e ‘_ Q . | , \ . _ -. . . . .1 . _- -- - i .. . -_ - . _ - - . .- .- _‘ . - I O , . ‘ t - 1 7 4 - ' . . > - . , . o . .. _ 1.:v ‘4. - v ‘ ' ‘ I . . _ _ ' . ‘ , v s .. ~ . . ~ . i .A‘ ' . .. . . . . . ~ . . .L 0, . , __ o s , , a. _~ . I . k I , . . . , _ I l ‘7 , ,_ ' . ~ . v . l . e - 4"- vs. ‘ ' w 1 . I .h ’ ‘ n t J ‘kl .1 4 u . ‘ .4 1 . -44- TABLE VII-b (continued) number of 3:1 F3 a... 19 25 giving (F ) colored non-colored Plats seedlings seedlings No. (ACr) (aCR 4 AcR) 57900 0 1 58000 2 2 58200 1 3 58300 2 1 58400 2 6 58500 2 2 58600 2 6 58700 1 3 58800 2 2 58900 1 3 59000 1 2 59100 0 0 2 59200 J __3 Actual total - 63 123 Expected total (1:2)- 62 124 Deviation +1 --1 M43LLQ=0.2 P.E. 4.2 ~45- Results of crosses between F“ A—, C-, and R- tester types and Emerson's.L-, -, and R-testers. Out of 394 3:1 F3 ears in the seedling test, 190 were selected for the 1926 planting. From such ears only seeds with yellow endosperm were planted in the field. 0f the 190 ears, 110 whose cobs were white and whose seedlings were non-colored were planted in 110 plate on one side of the field; while 13 red-cobbed ears with colored seedlings were planted in 13 plate, 25 red—cobbed ears with non-colored seed- 1ings were planted in 25 plats, and 42 white-cobbed ears with colored seedlings were planted in 42 plate on the other side of the field. The purpose of this planting was two-fold: (l) to identify the.L- ‘ tester type by crossing with Emerson's testers, and (2) to determine any relation between the color of seedlings and these three tester types, especially the A-tester type. The center rows in the field were planted with.Emerson's‘A-, C-, and R-testers. In each of the 190 plats, 4 or more plants were crossed with the Aptester, 4 with the C-tester and 4 with the R-tester. Several of the remaining plants in the plat were selfed. The principle for these crosses is shown in the following diagram: .Emerscnfe Aleurone color testers Tester types ‘A-tester C-tester R-tester from 3:1 F3 ears aCR AcR ,ACr aCR Colorless Colored Colored AcR Colored Colorless Colored ‘ACr Colored Colored Colorless v‘ . ' ‘ . s 4 ' e ‘ . .- . _ s , ' , ‘ - A' V, a . t . _ o 0 . . h . A - . * , I ~ .. . \ o I ‘ . - y a - - — _ u e x- . . v- e -46- In this way, the.A-, C-, and R-tester types from the yellow seeds of these 3:1 F3 ears could be determined and distinguished. There were 92. plats in which the plants or ears proved to be the A-tester types because in these plate 171 hybrid ears were color- less when crossed with the Emerson A—tester, 214 were colored when crossed with the C-tester, and 178‘were colored when crossed with the R—tester. The results are arranged in Table VIII-a and illustrated in Plate I. Some of the selfed ears in these plats are the strains of Aptester Yellow Dent sought for in this problem. .4 detail for selection of these strains will be given under another heading. Table VIII-b shows that plants or ears of 31 plate proved to be the C-tester types since they gave 53 colored hybrid ears when crossed with A-testers, 49 colorless ears when crossed with C-testers, and 4? partially colored and all colored ears when crossed with R— testers. The types are illustrated in Plate II. The latter hybrid ears should be all colored according to theory, as indicated in the diagram. The partially colored ears were probably caused by acme of the R-testers, being heterozygous for the 0 factor. Fifty-fine plats were found to have plants of the R-tester type as shown by the fact that the plants gave 8% colored ears when crossed with.A-testers, 7c colored ears when crossed with C-testers and 73 colorless ears when crossed with.R~testers. The data are indicated in Table VIII-c and types illustrated in.Idate III. There were 12 plate in which both crease and selfings failed. They are noted in parenthesis ( ) in Tables Ixsa and IXpb. Some of '0 “-47- TABLE VIII-a. Showing results of crossing the datester types (aCR) with Emerson's AP, C-, R-testers. name of (Access- F1 F2 13 Color No. ofeesNo. of ears variety ion No. 1923 1924 1925 1926 of F3 color- colored selec- selec- selec- Plat seed- less when crossed tion tion tion No. lings when with No. No. No. crossed with A-tester C-, R-tester Red Cobbed Duncan 234 31802 44807 52301 650400 w l 2 2 White Cobbed failed failed Duncan 235 301150 40701 52504 650500 w 0 4 0 40703 52702 650700 w 2 2 2 40709 53006 651200 w 3 3 3 301151 40802 53202 651300 w 3 4 3 53205 651400 w l 2 2 40803 53302 651600 w 3 1 3 40807 53404 651900 w 2 2 1 53405 652000 w l l 2 White Cobbed M.A.C. 254 30331 41102 53908 652400 w 3 4 3 53910 652500 w 2 4 3 53911 652600 w 2 2 1 53913 652700 w 3 4 2 53918 652900 w l 1 2 41109 54006 653000 w 4 4 3 54009 653100 w 1 3 1 41113 54103 653300 w 4 5 2 54112 653400 w 2 2 l w : non-colored (green) - — - fl - - - - - - - .- - - - - - a. - - - - - — - - — - - C. - fl - - - - - - - — - . flame of Access- F1 variety ion No. 1923 selec— tion NO. White Cobbed 254 M.L.C. 30332 30333 30408 -48- TABLE VIII-a. (continued) F2 1924 selec- tion NO. 41118 41119 41211 41212 41216 41311 41312 41401 41409 F3 1925 selec- tion No. 54405 54408 54505 54608 54708 54905 54906 54908 55005 55007 55103 55105 55303 1926 Plat no. 653500 653600 654000 654100 654200 654300 654400 654500 654800 654900 655000 655100 655200 655300 655500 655600 656000 656400 656500 656600 656700 Color seed- lings ITO. of color- less rNOe Of colored ears when ears when crossed crossed with with .A-tester C-, R-tester 3 2 1 3 4 2 3 2 0 2 2 3 3 3 2 3 2 1 0 0 2 1 2 1 2 3 2 2 0 1 3 3 3 3 3 3 4 3 3 3 (‘3 N ~49- TABLE VIII—a (continued) .Nsme of Access- ,F1 (F2 '33 Variety ion No. 1923 1924 1925 .1926 ' COIOI‘ INC. or ‘NO. or .selec- selec- selec- Plat of F3 color- colored ears No. No. No. lings ears when crossed crossed with with A-tester C—, R-tester White Cobbed 254 41413 55502 656600 w 1 2 l M.A.C. 55508 656900 w 1 3 1 55509 657000 w 3 2 2 55510 657100 w 2 2 1 55511 657200 w 2 4 3 30409 41503 55603 657300 w 1 2 2 41504 55703 657400 w 2 4 1 41507 55802 657500 w 1 2 2 55803 657600 w 3 4 2 41512 55903 657800 w 2 3 2 55904 657900 w 2 2 3 55909 656000 w 2 2 1 30513 41602 56007 658100 w 2 3 3 41605 56202 656200 w 2 2 3 30514 41701 56303 656300 w 2 1 0 41708 56503 658600 w 3 2 3 30515 41816 56801 658700 w 2 2 2 56802 656600 w 1 3 3 30604 41902 56902 659000 w l 1 4 56903 659100 w 1 3 2 41904 57003 659200 w 2 3 2 _ ' I _ .— ‘ .1 . -, 7 - -. -.. . ’ v . . 4 V ‘ ‘ 4 V , p .. ... t A . ‘7 ‘ . . ‘ . ‘ . , I i i. ) ' . 4 -‘ I A . l ' . . . ‘ . ~ . , ‘ . , . . 1 i .. ‘ 7 I ._ . i ‘7 . v . . . . . . D ‘ ‘ . . . -50.. TAfiLE VIII-s (continued) Name of .Access- “F1 .33 'F3 .1926 .Color .30. of .Ho. 0: 'Vsriety ' ion 1923 1924 1925 Plat of F3 color- colored ears No. selec- selec— selec- No. seed- less when tion tion tion lings ears when crossed No. No. No. crossed with with A—tester C-, R—tester White Cobbed 254 41905 57104 659300 w 0 2 1 M50- 30904 42401 57501 659500 0 31008 42501 57602 659600 3 42505 57703 659800 4 42516 57802 659900 2 31009 42602 57901 660000 2 1011 42701 58004 660100 3 42706 58103 660200 2 58107 660300 3 58108 660400 3 31111 42802 58202 660500 3 31112 42906 58303 660700 2 31113 43004 56403 660900 1 58404 661000 1 31306 43603 58501 661100 6 58502 661200 2 31308 43803 58602 661300 3 58603 661400 2 56606 661500 4 58607 661600 2 31309 43916 58702 661800 3 ‘Rame of Access- Variety ion No. Nelson X 255 Bailey Bailey 128 White Cobbed 235 Duncan Red Cobbed Duncan White Cobbed 254 MOAOCO 234 $1 1923 selec- tion NO . 32201 32301 32407 30806 31803 30331 -51- 'TABLE VIII-a (continued) .F2 1924 selec- tion NO. 45502 45702 46006 42203 44807 41118 'F3 1925 selec- tion NO. 59205 59206 51201 51402 51403 51502 57303 57305 57403 52302 54312 1926 Plat No. 662000 662100 663500 663900 664000 664300 665500 665600 665700 666400 667900 Color Of F3 seed— lings TOTAL . . . . . . .'. . . . . . 9€Lplats . . 'NO. Of 1R0. Of color- colored ears less when ears when crossed crossed with with ‘A-tester C-, R—tester ..ii' 171 1 2 3 2 1 1 1 1 2 1 1 1 o o o o 1 o 2 1 ._JL ._Ji 4 216% 1ma 4Q HAN! Individuals in plat 652400 proved to be the A-tester type (11011) of corn since they produced three colorless ears (tsp of the picture) when crossed with'herson'e A-tester, three colored ears (on left side at bottom) when crossed with G—tester and two colored ears (on right side at bottom) when c}cssed with R-tester. e"\ -52.. $4813 VIII-b. Showing results of crossing the C-tester types (AcR) with Emerson's A-, 0-, and R- testers. Name Access- 7F1 F3 F3 - 1926' Color ‘ E23062 of ears or 103 1933 1934 1925 Plat 0f F3 'colored colorless colored variety No. selec- selec- selec- No. seed- when when when tion tion tion lings crossed crossed crossed No. No. No. with. with with .A-tester C-tester R-tester Btiley 128 32201 45502 51204 650000 w 2 3 3‘ White Cob. 235 301150 40703 52703 650800 w 1 1 0 hmcan , 10702 52603 650600 w z 1 1“ 40705 52803 650900 w 2 2 3‘ 40709 53002 651100 w 2 5 3‘ 301151 40803 53303 651700 w 2 2 1 + 1‘ 53304 651800 w 2 2 3‘ White Gob. 254 30255 40901 53502 652100 w 1 1 2‘ M400. 40902 53601 652200 w' 3 2 2‘ 30256 41002 53803 652300 w 4 5 1 4 2‘ 30333 41313 55202 656100 w' 5 2 1‘ 30409 41507 55804 657700 ‘W 1 ' 4 2‘ 30514 41706 56401 658400 w 2 2 3‘ 56403 658500 w 3 O 3‘ 30515 41816 56804 658900 w 2 2 1‘ 31008 42501 57603 659700 w 0 2 1‘ 31111 42802 58204 660600 w 0 1 0 31309 43916 58701 661700 w 2 1 2 53104 66(900 w 3 3 1“ Bailey 128 32201 45502 51202 663600 w 1 0 0 51205 663700 w 2 1 2‘ 45509 51319 663800 w 1 1 1 32301 45702 51405 664100 w 1 2 1 51406 664200 W 1 1 0 - g _. O . _ .. . L .- ._ ._ _. _ _ a -- _ —. _ _- — -- _ - , -. . . ' . ' ~ - ' ’ \e , . ' e . t , . ‘ < e _.. ‘ _ ‘ ' . . u .- s 1 - t. , I V ' A I . e _ o 7 - v . .4 .. - . - . . . . ‘ A I , l (I v . ' . - . ‘ . ' n r 1 . . , -- - -‘ . ‘ , 1 . _ p v 1 h, ' . - -53- TABIE VIII-b (continued) flame of Access- 31 32 ,F2 1926 . Color E number 9; gen Variety ion No. 1923 1924 1925 Plat of F3 colored colorless colored selec- selec- selec- No. seed- ‘ when when when tion tion tion lings crossed crossed crossed No. No. No. with with with A—tester G-tester R-tester Red Gob. 234 31604 44306 51701 664400 w 2 0 2‘ Duncan 61801 664500 w 0 0 1 44311 51802 664600 w 1 0 0 31704 44609 52003 664800 w _ l O 1‘ 31802 44711 52201 665100 w l 1 0 31430 44104 59102 665900 w 2 1 3‘ 31432 44207 59 204 666000 w _3_ _1_ __L‘ TOTAL. . . . . . . . . . . 3f plats . . . 53 4D ‘48 ‘ hrs were partly colored. N . - ‘ s O _ _ u o n e x 1 O o O . n , ,. _ . H , ., O . n u .. ,. _ . ~ e _ , e I .. e . . 1 . u ”H u . . h .(C _ 1 .. a .l ‘ mm 11 FA 137f fine picture shows that individuals in plat 652300 proved to be the Gétester type (LcR), since they gave three colorless ears (center or the picture) when crossed with Emerson's O-tester (AcR), three colored ears (on left-hand side) when crossed With A—tester and two partly colored ears (on right-hand side) when crossed with Rptester. Ihe partial coloring was probably due to heterozygous {00; in_RFtester. -54— ‘TABLE VIII-c. Results of crossing the R—tester types (Aer) with Emerson's A-, c-. and R-testers. flame of Maternal 3‘], P2 F3 1926 Color Variety Aooosa- 1923 1924 1925 Plat Ioi’ 33 colored when colorless ion N0. selec- 86160- 86160- 'No. seed- crossed with when crossed tion tion tion lings with with No. No. No. A—, 0- tester B-tester Bailey 128 32408 46006 51503 650100 B 1 2 1 Red Cobbed 234 31604 44306 51704 650200 R 3 1 0 Duncan 51707 650300 R 3 3 2 White 00b. 235 301150 40708 52902 651000 R 1 1 1 Duncan White Cob. 254 30331 41102 53917 652800 R 2 2 4 MoA.0. 41113 54103 653200 R 1 2 1 542.08 653900 R 1 1 0 30332 41211 54604 654700 R 2 3 1 30333 41311 55004 655400 R 2 2 2 55008 655700 B O 1 0 55010 655800 R 2 1 2 41313 55203 656200 R 3 3 3 55204 656300 R 2 2 3 31306 43603 58402 660800 R 2 3 2 Bailey 128 32201 45502 51203 662200 R 3 2 2 45509 51315 662300 R 2 1 1 51317 662400 11 1 0 0 32301 45702 51404 662500 R 2 1 1 51413 662600 R 1 3 3 Red Cobbed 234 31604 44304 51601 662700 R 2 2 1 Dmcsn mmber 9f ears R 3 colored 2’ -55- 'TABLE VIII-c (continued) he of Maternal F1 32 ,F3 1926 -Color vMbe: 9: car: Viriety .Lccess- 1923 1924 1925 Plat of F3 .colored when colorless ion No. selec- selec- selec— No. seed— crossed when crossed tion tion tionn lings with with N0. No. No. . Ag, G-tester R-tester 44303 51602. 662800 R 2 1 1 51603 662900 R 2 l 1 44306 51703 663000 R 2 2 1 31703 44503 51904 663100 R. 2 l 2 51905 663200 R 1 0 2 31803 44807 52303 663300 B. 1 1 1 White Cobbed 264 30806 42203 57306 663400 R 2 l 0 ”‘00. Red Cobbed 234 31802 44711 52205 665300 R 1 1 2 Duncan Bailey 128 32201 45502 51316 666100 R 1 2 1 Red Cobbed 234 31604 44306 51702 666200 R 1 1 1 Duncan 31704 44609 52001 666300 B. 3 1 0 (White Cobbed 235 501150 40701 52507 666500 R 2 1 2 Duncen 52510 666600 R 0 0 2 40710 53101 666800 R 1 0 2 301151 40803 53305 660000 73 l 2 2 40807 53403 667100 R 1 0 2 White Cobbed 254 30331 41102 53904 557300 R 1 1 0 “0‘00. 53905 557300 R 2 1 2 53906 557400 R 2 1 2 55913 557500 R 2 1 2 41109 54005 667600 R 2 1 1 41113 54111 667700 11 1 z 2 -55- 'EABLE VIII-c (continued) .— ‘Namc of Maternal F1 'F2 'F3 1926 'Color Viriety .Access- 1923 1924 1925 Plat of F3 colored when colorless ion N0. selec- selec- selec- N0. seed- crossed when crossed tion tion tion lings with with N0. N0. No. .A—, C-tester R-tester 41114 54204 667800 R 1 1 2 54205 653700 R 0 1 0 41119 54406 668000 R 2 2 1 30332 41201 54504 668100 R 1 1 1 30333 41212 54704 668200 B. 1 l 1 41312 55104 668300 R l O 0 30408 41409 55412 668400 R 0 0 1 30515 41802 56604 668500 R 2 1 2 30806 42203 5 7409 665800 R 3 0 0 31111 42906 58302 668600 R 2 2 1 31306 43603 58504 668700 R 2 2 1 Nelson 1 255 31430 44004 58905 668800 R 1 1 1 Bailey 31432 44207 59202 668900 R 1 ' 1. 1 TOTAL e e e e e e e e e e e e e e 55p13t8 e 88 71 73 'PLLTE III ‘a. Mo 5 '-, Koo Individuals in this plat proved to be the R—tester type (LGr) as they produced two colorless ears (right) when crossed with R-tester, three colored ears (center) when crossed with C-tester and two colored ears (left) when crossed with A—tester. -57.. them were determined as A—, or R-tester types by their seedling color. Glancing at Tables Ill-a and IX—b, one will see that there is perfect correlation between the R-tester type and the color of seed- lings. In other words, those whose seedlings were colored were the B-tester types. Likewise, the A-tester types were correlated with non-colored seedlings, but they could not be distinguished by seed- ling color from the C-tester types whose seedlings were also non— colored, when both types came from the 27:57 F2 group (see Table Iii-b). Thus it is seen that these data give a substantial support to the seedling test. DESCRIPTION AIID SELECTIOII OF STRAINS 0)." 5323mm YELLOW I) ‘m com: In 92 plats a total of 371 selfed ears proving to be the A- tester types was obtained in the F4 generation (1926) . Arnong these, some ears were yellow dent and some were yellow flint. In regard to the endosperm color, some ears were homozygous for the yellow endosperm and some were segregating for the yellow and white endo- sperms (see Table VI). The flint and the white endosperm characters came from Emerson's A-tester, while the dent and the yellow endOSperm characters came from the maternal strains used in this escperiment. Out of the 371 ears, 36 with homozygous yellow endosperm in dent form were considered as desirable ears. The pedigrees of these ears are given in Table 3.. r‘rom the 36 ears, 11 were selected as --58- ~QABLE pra. Showing that the seedling color of colorless ,individuals of 3:1 F3 ears coming from the 9:7 5‘ group were verified 2 by crossing with Emerson's testers. The Antester type resulted from those individuals having non-colored seedlings and the B—tester type from those having colored seedlings. 1925 'Tester R : colored; W = non-colored or 1925 ' o ' ”3’ minus (41 type )2? sailing i??? T2533” t:::°§;. color Pigt Ezozgd tiselec- color Plfit proved . - on No. No. to be- 51501 R 552700 Lsr 54112 71 553400 aCR 51502 R 552500 a 54115 7: 553500 " 51503 E 552900 '5 54115 17 553500 " 53403 s 557100 '5 54204 1: 557500 ACr 53404 3 551900 505 54205 5 553700 ( N ) 53405 :7 552000 9 54207 R 553500 ( n ) 53904 E 557200 40:- 54205 14 553900 " 53905 5 557300 " 54304 17 554000 5011 53905 R 557400 '3 54305 3 554100 .. 53905 27 552400 5011 54310 w 554200 " 53910 :7 552500 n 54312 17 557900 N 53911 17 552500 '1 54404 17 554300 " 53913 s 557500 151. 54405 17 554400 n 53915 5 552700 303 54405 5 555000 Aer 53917 1 552500 AC: 54405 11 554500 402 53915 3' 552900 aim 54504 5 555100 ACr 54005 5 557500 Acr 54509 :7 554500 (505) 54005 s 553000 5011 54504 R 554700 ACr 54009 71 553100 " 54505 :7 554500 505 54103 s 553200 ACr 54505 11 554900 v 54107 17 553300 aCR 54704 1‘: 555200 40:» 54111 R 557700 ACr 54705 " 555000 505 -59- EABLE pra (continued) 1925 13 1926 Tester 1925 (F3) 1926 Tester (33) seed— (3;) type (F3) 0010; of (F4) type Selec- ling Plat proved Selec— Plat proved tion color No. to be- tion Seed- No. to be- No. No. ling 54905 W 655100 aGB 56604 .3 668500 ACr 54906 W 655300 ” 57005 W 659200 aCR 54908 W 655300 * 57203 W 659400 ( " ) 55004 E 655400 ACr 57303 W 665500 ” 55005 W 655500 80R 57305 W 665600 ” 55007 W 655600 " 57306 R 663400 ACr 55008 E 655700 ACr 57405 W 665700 803 55010 R 655800 “ 57404 R 665800 ACr 55411 5 555700 555 57703 5 559500 505 55412 E 668400 ACr 58103 ‘W 660200 " 55502 W 656800 aCR. 58107 W 660300 “ 55508 W 656900 " 58108 W 660400 ” 55509 W 657000 “ 55510 N 657100 " 55511 5 557200 9 ( ) crosses in these plats failed, 55603 W 657300 " so types were detennined by seedling 55903 W 657800 " 3 color. 55904 W 657900 " 55909 W 658000 " 56007 W 658100 u 66202 5 555200 5 56303 3 555300 5 56503 W 658600 " EARLS LK~b. Snowing that the seedling color of the color- less individuals of 3:1 F3 ears coming from the 27:37 52 group were verified by crossing with Emerson's hp, C-, and R-testers. The R-tester type was correlated with colored seedlings; both .A-, and C-tester types resulted from those having non-colored seedlings. 1925 F3 1926 Tester 1925 F3 1926 Tester type (F3) seed- (Eh) type (F3) seed- (3;) proved Selec- ling Plat proved Selec- ling Plat to be- tion color No. to be- tion color No. lb. No. 51201 W 663500 GCR 51707 R 650300 .ACr 51202 W 663600 AGE 51801 W 664500 AGE 51203 E 662200 .Adr 51802 W 664600 ” 51204 W 650000 .403 51904 E. 663100 ACr 51205 W 663700 " 51905 E 663200 ” 51315 R 552300 ACr 52001 R 555300 " 51316 R 666100 ” 52002 W 664700 (-—) 51317 _ 11 552400 1' 52003 :7 554500 453 51319 W 663800 AcR 52101 W 664900 (--) 51402 W 663900 303 52201 W 665000 (4:) 51403 W 664000 ” 52202 XI 665100 A03 51404 E 662500 ACr 52204 W 665200 (--) 51405 W 664100 7103 52205 B 665300 ACr 51406 W 664200 " 52301 W 650400 803 51413 .R 662600 7ACr 52302 W 666400 " 51502 W 664300 208 52303 R 663300 7ACr 51503 E 650100 ACr 52401 n 665400 f--) 51701 3 554400 155 52504 (7 550500 502 51702 E 666200 .LCr 52507 3 666500 (ACr 51703 R 663000 " 52510 R 666600 " 51704 E 550200 '5 52503 77 550500 155 colored; W = non-colored -61.. EABLE LXpb (continued) 1925 F3 1926 Tester 1925 13 1926 Tester type (F3) seed- (EA) type (E3) seed- (5;) proved Seleo- ling Plat proved Selec- ling Plat to be— tion color No. to 115- tion color 110. No. No. 52702 71 550700 555 55205 R 555200 ' ACr 52705 71 550500 455 55204 5 555500 N 52505 550900 " 55205 555400 5511 52902 551000 ACr 55505 555500 " 55002 551100 Add 55507 555500 N 55005 555700 (ACr) 55705 557400 5011 55005 551200 5511 55502 557500 N. 55101 555500 ACr 55505 557500 9.511 55202 551500 5011 55504 557700 155 55204 555900 (ACr) 55401 555400 .41. 55205 551400 5511 55405 555500 .. 55207 551500 (--) 55501 655700 am 55502 551500 5011 55502 555500 " 55505 551700 11511 55504 555900 .452 55504 551500 '- 55902 559000 801 55505 557000 Aer 55905 559100 " 55502 552100 5511 57104 559500 11 55501 552200 " 57501 559500 " 55505 552500 1' 57502 559500 '5 55102 555900 (ACr) 57505 559700 1511 55105 555000 555 57502 559900 555 55104 555500 Aer 57901 550000 *7 55202 656100 .108 58004 660100 n -62- TABLE IX—b (continued) 1925 p 1925 Tester ‘ (B ) 3 3 seed- (F4) type Selec- ling Plat proved tion color No. to be- NO. 58202 ‘7'} 660500 20.11 52002 58204 W 660600 £03 52101 These individuals 58303 W 660700 803 52204 , could not be classified _ 5830?. Id 668600 101‘ 52.401 1 as the A-, or C-tester 58402 E 660800 " 53207 type by seedling color. 55405 '5 550900 555 52501 1' 55404 1'! 551000 " 55501 5' 551100 9' 5;}; 58502 W 661200 203 } were of the ACr type 55102 58504 R 668700 ACr as determined by their 58602 W 661300 803 colored seedlings as 55505 5 551400 " crosses 551155 in the 55505 17 551500 '5 r15 ld . 58607 W 661600 " \' 58701 ‘3 661700 103 58702 W 661800 2011 58704 W 661900 ‘03 58905 R 668800 101' 59102 W 665900 45.5 59202 E 668900 ACr 59204 17 666000 10R 59205 W 662000 5011 59206 W 662.100 “ (--) crosses in these plats failed. -53- the most desirable types and are sheen in Table K.with (*) marks and in Plate IV. This table shows that one of the 56 ears is the descendant 434 of White Cobbed Duncan, :5 are descendants of Emits Cobbed Golden Glow .x White Cobbed Duncan, Ears numbered 656510 and 656511 are the A—tester yellow dent with.white caps, and hence are called the "White Capped A-Tester Yellow Dent". They are sisters as both came from a mother ear (NO. 55303) of the F5 generation. ‘15 to some ear characters, all of the 36 cars are white cobbed with yellow endOSperm in the dent form: the number of kernel rows ranges from 10 to 14; the length of ear ranges from 10 to 19 centi- meters; the weight of ear is from 34 to 126 grams; and the indentation of all ears is smooth. The growth, as observed in the field in 1926, of.A-, C-, and R—tester types was fairly good, although they were hit by a sand storm and several frosts during the seedling stage. The soil was very poor because of lack of manure and fertilizers, but plants of the.A-teSter type in most plats looked more uniform and vigorous than those of their sister types, C-, and R-testers in the same breeding lot (see the Table on pagebo). This gives at least an indication that the lack of anthocyanic pigment in the A-tester types has no bad effect upon the plant growth and the yield. It was thought that the.L-tester types, lacking the factor for anthccyanin, would grow weaker and not give as good a yield as . s a U . _ . u .. . . . . . . . V a . 1 I a 1 ) l . '5 . .\ ,-54_ TABLE X. Showing parentages and general characters of 56 strains of.A-tester Yellow Dent. Name of .Aocess- _P1 F1 F2 F5 E4 ch Length No. Wt. Color Indenta- Variety ion No. 1922 1923 1924 1925 1926 Color of ear of of of tion selec- selec- selec-selec- selec- in rows ear endo- tion tion tion tion tion cms. in sperm grams Wh, Cob. 235 02600 501150 40803 55302 651608 White 13 12 68 Yellow Smooth Duncan 55.050. 254 03599, 50551 55905 552411* N 15 14 97 deep N G.Glow 12904 yellow 11 552412 N 14 14 97 yellow N Wh.ch. Duncan 4110 55910 .552510 N 15 14 55 N N .55915 1552905 N 12 14 59 N N ,555012N N 17 12 110 N N {553013. N 15 14 101 N N 41109 54005f555014 N 15 14 55 N N 1555015 N 14 14 55 N N (555015 N 17 12 100 N N 41115 54105 555512 N 19 14 55 N N 54010. N 14 14 95 N N 554011N N 15 12 77 N N ,554012 N 15 12 75 N N 41115 5450 554015 N 12 ‘ 12 70 N N 554014 N 12 12 55 N N ,554015 N 12 12 50 N N 51,. . 5 . 25202 50405 41211 54505 554505 N ~ 10 12 54 N N J 554507 N 12 14 55 N . N 55505 555510* N 15 12 105 wh.5ap. N yellow 41401 656511‘ " 13 12 96 " " (‘) The most desirable strains. "73 u-O‘J- TAB“: 1’1 ( co nti nue d) 'Name offiAcoess— P; ‘F1 F2 variety ion 1922 1925 1924 No. selec- seleo- selec— tion tion tion No. No. No. 41409 .'= . - ‘ 25206 31008 42416 31009 42602 ,42701 31011 42708 25207 31111 42802 25313 31306 43603 31309 43916 F3 1925 selec- tion NO 0 55307 55411 57802 57901 58004 5810? F4. 'Cob'Length Ho. 1926 Color of of seleo- ear rows tion in No. ems. 555510 Ehite 15 12 555707 N 10 14 559907* N 14 12 559905* N 15 12 550005 N -- -- 550110 N -- -- 550209 N 15 12 550210 N 12 14 550505 N 17 14 550509 N 11 14 550505 N l7 10 551112 N 16 12 551205 N 12 14 551507* N l3 14 551505* N 12 12 ‘Ut. ‘Color 'Indent; of of tion ear endo- in sperm grams 108 wh.cap. Smooth Yellow 45 Yellow " 90 N I. 93 H H __ a n __ n u 68 faded " yellow 74 yellow & " dilute yellow 126 yellow " 34: N l' 89 0 ll 9 l it It 75 t! I! 109 deep " yellow 9 5 N M t . \ Strains of A-tester Yellow Nos. 656510 and 656511 (on right side the White Capped A-teeter Yellow Dent. Dent Corn. or the picture) are -55- the C-tester and R-tester types, which have the A factor. In order to put this discussion on a scientific basis, a statistical study of plant height, ear length, and ear weight for these three tester types, A-, 0., and Ba, was made. Twenty-five plants were measured for height if the plat had 25 or more plants, and all plants were measured if the plat had less than 25 plants. All selfed ears were measured for length of ear, and weight of ear. Biometric constants were calculated. A cmnparison of height of plants was made between the A-tester type and the R-testsr type. The difference was slightly in favor of the A—tester type, but it was not significant statistically. Comparisons of ear length and ear weight between these two types indicate that the differences are also in favor of the A-tester type, but of not statisti- cal significance. Similarly, these three variables hsve been compared between the A-tester type and the G-tester type. All differences were in favor of the A-tester type. ‘Ihs data for these comparisons are given below Type ' Height of plant Length of Bar Weight of Ear Mean ‘ B C.Y. Mean k E 0.7. Mean * B 0.7. Feet Percent (32g. Percent Grams Perm A-toster 4.98 g 0.38 11.56 13.41 h 1.13 12.83 64.09 3 9.54 22.20 R—tester 5.26 g 9.32 11.33 83 12 3 43 52,22 3 2.20 28.33 D11.-." +0.02 1 0.53 +0.23 +0.58 :2 1.56 —0.60 412.87 $13.23 —6.12 ‘-t68ter 4098 t 0038 11056 13e41 * 1.13 12.83 54009 i 9054 22020 G-tester 5.2820,57 15,65 12.044.11.18 14,20 51:61:19.0; 29,27 Dif.--— 0.00=t0.64 -3.10 +1.37gl.l4 -l.87 +12.484-13.79 -7.07 Summing up the above, it is concluded that the A-tester type of corn is not inferior to the other types, C—, and R—testers, in growth and yield, in terms of plant height, ear length, and ear weight. a. _ 7 . _ .. . , i l l ’ . . .. § I O . . . . . A - I . \ .— .- 0 , . . ‘ \ -- .. - . .- -6 7... RELATION 05 11115 540220115 113 *0) rr 1‘0 'l‘iE 572102113115 0F.ANTHOCIANIC PIGHENT AS FOUND IN THIS EXPERIUENT Emerson (1921) has pointed out that the R factor exists as a series of allelomorphs, such as, Br, 85, rr, r5, etc. The factor rr is recessive for aleurone color and dominant for plant color, while the factor 38 is dominant for aleurone color and recessive for plant color. RrRr or Brag or rrrr or rrr8 will cause more or less red pigment to develop in the plant body in the presence of the A.factor. Neither R326 nor rgrg will cause red pigment to develOp in the plant body in presence 02.4, which is the factor reaponsible for the produc- tion of any anthocyanic pigment in a corn plant. It has been.mentioned in a previous section that the.A-, and C-tester types, progenies of the 27:37 F2 group, which could not be separated by the seedling test, might involve the factor R585, and that the B-tester types, which were easily rogued out by seedling tests, must involve the factor rrrr. It is upon this fact and upon this assumption that the following investigation has been taken up. Seedling color in the P1 and 31 generations Remnants of 16 maternal P1 strains were obtained and planted in sand trays. TWelve of them gave only colored seedlings, and the re- maining 4 strains gave mostly colored seedlings with but very few non— colored ones. This indicates that the strains whose seedlings were pigmented must have carried the factor rr which causes the seedlings to be colored in the presence of the.A factor. Their compositions -68- TABLE XI-e. Showing that all maternal P strains gave colored seedlings, indicating the presenoeu of rrrr. Name of Access- 1922 1923 number of seedlings variety ion No. Selection Plat No. No. or Colored Non-colored Plat No. White Cob. 93690 0. Glow x 12901 30200 22 0 White Cob. 254 22599 3 Duncan 12904 30300 9 4 25202 30400 19 0 25203 30500 22 0 25303 30600 16 0 25306 30700 21 0 25312 30800 22 0 25205 30900 22 0 25206 31000 35 0 25207 31100 34 2 25302 31200 38 0 Nelson X 255 23000 31400 16 0 Bailey Red Cobbed 234 234001 31600 15 5 Duncan 23701 31800 25 14 23802 31900 5 0 Bailey 128 24301 32300 20 3 TOTKL --------------- 371 - - - 28 TABLE XI-b. The paternal parent, A-tester gave all non-colored seedlings. These were later proved to be RSRg. AEar No.— -Colored seedlings- -Colorless seedlings- 1 0 24 2 0 22 3 0 24 4 _9 25 mom ------- o ————————— E PEATE V This picture shows that the maternal Pl strains give colored seedlings, indicating the presence of rrrr. -69- might be written as AACC rrrr, AACcrrrr and AAccrrrr. The results are in Table XI-a and illustrated in Plate V. Fbur ears of Emerson's A-tester corn which were selfed in 1926 were planted in sand boxes and produced all non-colored seedlings. The data will be found in Table XI-b. The composition of the Emerson.A-tester was either aaCCRgfig, or aaCCRyhg, or aaCCRrhf Assuming the first type to be the case, in the £1 generation, the seedlings from any of the following crosses should be colored be- cause rr is dominant over RS. -Materna1 strains- -Emerson's‘A-tester- AACCrrrr x aaCCRgag AACcrrrr X aaCCRShg AAccrrrr .K AACChghg ‘Unfortunately, there were no remnants of F1 seeds left, so there is no way to get data for this generation. This supposition therefore, remains to be proved by the seedling behavior in the later generations. Seedling behavior in the FE and 35 generations Remnants of 66 F2 ears used for planting in 1925 were tested. They were colored kernels from both 9:7 ears and 27:37 ears. All were planted in sand boxes in the laboratory. If the hypothesis at hand is right, the FE colored seeds either from the 9:7 ears or from the 27:37 ears must give colored and non-colored seedlings in the proportion of 2 to 1 according to the following factorial diagrams in which the EB behaviors are also shown. . - n ‘ , 4 - . \, I . v , s i o- , o - , ' u o . o . ‘ n I . ' s.(‘ F . _ , o a ., s 979— Diagram a. Colored seeds of 937 E2 ascer’rr X aaccngsg. Genotypic 12 genotypes PrOport Ratio 1 AACCRQRg giving non-colored seedlings 2 AaCCRgRg giving " " 2 AACCRgrr giving colored seedlings 4 £30038ng giving colored seedlings Diagram b. Colored seeds of 27:37 AAccrrrr X aaCCligl'tg. Genotypic F2 genotypes Pr0portion of Ratio seedling color 1 A1003i f non-colored seedlings ] 2 AaCCBgRg n n n ! g g ‘ S 1 2 MCCR ll " " " I ‘ l 4 “503839; I. u u j 2 AACCRgrr colored seedlings 4 AaCCRgrr .. N z 4 uCcRgrr " " 7 2 a a .AaCcRgrr .. j ,2 “303 it ears from F 3 behavior ion of seedling color breeds true for non- colored seedlings l LAOGR§BS all non- 2 ABCCRERS colored l aaCC seedlings 1 uccsgng colored 2 AACGRBrr seeds giving 2:1 of color- ed and non— colored seed- lings Colorless seeds giving only colored seedlings 2 colored; 1 non- colored seedlings as in FE l AACCrrrr F2 ears from £3 seedling behavior Breeds true for non-colored seedlings 1 AACCRéRg all non-colored 2 AaCCRSflg seedlings 1 saCGRglig l AACCRERE all non-colored ' seedlings l AAccRbflg .— o- l MC-Ciiglig non—colored Seedh'ns 2 AACCRgrr colored lAACCrrrr all colored 2:1 colored and non-colored -88 111 F2 3:1 colored and non-colored as in E3 2:1 colored and non-colored as in F2 ll -71- TABLE XII. Colored seeds from 9:? EB ears and 27:57 En ears gave colored and non—colored seedlings in a ratio of :1, thus indicating the segregation of the allemorphic pair Rérr. name of Access- 1923 1924 Ratio Number of seedlings- Variety ion (3 ) (E2) of No. sel c- selec- ear Colored Non-colored tion tion ' NO. NO. White Cob. 255 301150 40701 27:37 16 6 Duncan 40702 " 6 6 40703 " 3 1 301150 40705 " 4 3 40708 “ 4 2 40710 " 13 7 301151 40802 " 6 6 40803 '1 o 5 40807 9:7 13 11 White Cob. 254 30255 40901 27:37 7 3 G.Glow _ K' 40902 " l 2 White 00b. . Duncan 30256 41002 " 15 7 30331 41102 9:? l7 6 41109 “ 19 7 4lll3 " 13 6 41114 '5 10 a 41118 '3 11 7 4lll9 " l2 8 50332 41201 " 7 4 41211 " 9 4 -72- TABLE XII (continued) flame of .Access- 1923 1924 Ratio Number of seedlings- Variety ion No. (Fl) (Eh) of selec- selec- ear Colored Non-colored tion No. tion No. 254 30332 41212 9:7 ‘ 12 4 (continued) 41213 27:37 4 1 41216 " 16 7 30333 41311 ” 0 18 41312 " 14 7 41313 " 1 0 30408 41401 9.7 2 1 41409 " 9 5 41413 e 12 7 30409 41503 e 15 6 41504 27:37 6 2 41507 " 5 4 30513 41602 9.7 14 6 41604 " 13 8 41605 a e 5 30514 41701 " 11 6 30515 41802 " 17 6 41810 " 16 9 41816 27:37 7 3 30604 41902 " 7 4 41905 9:7 15 8 30702 42002 " 8 4 -73.. TABLE XII (continued) name of iaccess- 1923 1924 Ratio number of seedlings Variety ion Ho. (Pl) (E2) of selec- selec- ear tion No. tion N0. Colored Non-colored _ - _ _ _ - - E5; .......................... --_ (continued) 30806 42203 9:7 9 5 42209 " 14 7 30904 42401 27:37 11 8 31008 42505 927 15 5 42516 27:57 3 1 31009 42602 " 7 2 31011 42701 27:37 17 6 42708 9:7 14 7 31111 42802 27.37 11 a 31112 42906 " 2 1 31113 43004 “ 8 3 31306 43603 “ 9 5 31308 43803 9:7 12 9 Nelson X 255 31430 44001 27:37 15 6 Bailey 31431 44101 " 5 2 44104 " 12 6 31432 44207 " 4 1 31604 44304 9:7 10 12 44306 27:37 3 0 44311 " 11 5 -74- TABLE XII (continued) Name of (Access- 1923 1924 Ratio number of seedlings variety ion No. (31) (Eb) of seleo- selec- ear Colored Non-colored tion No. tion No 255 31703 44503 27:37 3 2 (continued) 31704 44609 " 1 2 32201 45502 " 5 2 32301 45702 1' a 1 66 ears Actual total . . . 601 . . .-. 320 Expected tom]. (281). o 614 o o o o 307 D8V18510n o o o o -13 o o o 0N3 .2211:=.JIL. = 1,4 PoE. 9.6 Such was proved to be the case. Colored seeds of these 66 F2 remnant ears gave a total of 601 colored seedlings and 320 non-colored seedlings where 614 colored seedlings and 307 non—colored seedlings were eXpected. This exhibits a deviation of 15, which is only 1.4 times its probable error, indicating that observations were very close to expectancy. The results are arranged in Table XII.« ‘Unfortunately, many of the Rb ears had been thrown away before this seedling situation was fully realized. Those ears which remained were tested in two groups, i.e. those derived from.the colored in- dividuals of the 9:? F2 ears and those derived from the colored in- dividuals of the 27:37 Fé ears. F3 seedling behavior in the 9:7 Eb group In this group 6 F5 homozygous colored ears (1:0) gave only non- colored seedlings. The results are shown in Table XIII-a. Here is a critical test for the hypothesis indicated in diagram.a.’ These colored seeds of 1:0 F ears should carry the Rgfig factor in order to produce‘ 3 non-colored seedlings in presence of the.L factor. The Rghg'must have come from Emerson's.L-tester (aaCCBgflg) in order that the 1:0 Eb ears could have the constitutionaaccflgfig because it has been shown, Table XI-a, that rrrr came from the maternal P1 strains. Otherwise, these 1:0 Fa's, indicated in Table XIII-a, would not produce all non-colored seedlings. One hundred and nine 3:1 F3 ears showed all non-colored seed- lings either from the colored seeds or from the colorless seeds. These -76.. ears were proved1 to be the.A-tester types (a‘R in Table pra) with reapect to their colorless seeds. Consequently, they must have contained Rghg in order to produce all non-colored seedlings in presence of the.a factor. The data are set forth in Table XIII-b. Table XIII-c shows that another set of 97 3:1 F3 ears gave 1684 colored seedlings and 681 non-colored seedlings from the colored seeds and gave only 2083 colored seedlings from the colorless seeds. lExpected numbers of colored and non-colored seedlings for the former are 17102855 on a basis of 2:1. The deviation is 26, which is 1.6 times its probable error. This eXplains the fact that all these ears have been proved to be the R-tester type (ACr) with regard to their colorless seeds (see Table IXna), They came from the F2 genotype AACCBgrr (diagram a). Consequently, from such 3:1 F3 ears the seeds with AelCCIdgrr would give colored seedlings and the seeds with 8.4003515 would produce non—colored seedlings in the proportion of,2 to l. The colorless seeds with_AACCrrrr of the same ear would produce all colored seedlings on account of the presence of rr in addition to the A factor. Colored seeds from 43 9:7 F5 ears gave a total of 831 colored seedlings and 458 non-colored seedlings where 859.3:429.7 are expected 1 Not all of the 3:1 Eb ears were included in the 1926 planting. The ears which were selected for crossing with Emerson's testers were chosen because of characteristics other than those affected by Rg or rr and hence the ears reported in Tables IXna and IXpb may be considered to be random samples of the ears tested in the sand trays, and the results obtained by the field crosses may be considered as typical for the various types of seedlings. The lameholds true for the other results from 3:1 53 ears mentioned in this section. 7 I. TABLE XIII—a. Showing that only non-colored seedlings g were produced by the 1:0 F3 ears which.were progenies of the 9:7 FE group. { 1925 1 1 (s ) ‘ , 4 ~ 3 Number of seedlings : fl Selection [ No. Colored Non—colored 1 a : § 53901 0 18 V? 3 ; 53902 0 16 i 4 i ; 54101 0 14 i. 1 55401 0 s i' g p 56001 0 24 i 4 56002 _J; go 5; 1; Total 0 100 1‘ i E '1 i j': TABLE7XIII-b. Showing that Only non-colored seedlings resulted from both colored and colorless kernels of 109 3:1 F3 ears, 1925 (33) Selec- tion NO 0 From colored progenies of the 9:7 F2 group, indicating the presence of R532. From color- 1925 From colored From color- (35) ggzggls lag; gernelg Selec- -seedlings- -seedlings- tion colored oolor— colored color- No. less less 53404 0 24 0 25 53405 0 22 0 23 53908 0 20 0 11 53910 0 25 0 20 53911 0 23 0 18 53912 0 24 0 21 53915 0 23 0 22 53918 0 25 0 23 54004 0 24 0 21 54006 0 23 0 20 54007 O 21 O 15 54008 0 24 0 22 54009 0 24 0 21 54106 0 25 0 23 54107 0 24 O 25 54108 0 26 0 24 54110 0 21 0 23 54112 O 24 O 25 54116 0 21 0 20 54117 0 21 O 20 54118 0 21 0 15 54304 54305 54306 54308 54310 54312 54408 54404 54405 54.08 54512 54513 54514 54515 54605 54607 54608 54703 54705 3222212 -seedling§=_ colored color- less 0 20 0 15 0 20 0 25 O 21 0 23 0 25 0 20 0 19 0 20 0 19 0 20 o 25 0 24 0 14 0 20 0 21 0 24 0 20 0 25 0 21 less—kernglg -seedligg§- colored color- less 0 23 0 19 0 18 O 21 0 20 O 20 O 19 0 23 0 21 O 24 0 24 O 15 0 23 0 15 0 l3 0 24 0 21 0 25 0 15 0 20 0 15 ‘1925 (Pa) Selec- tion No. 54708 54905 54906 54908 54913 55005 55006 55007 55009 55405 55408 55409 55410 55411 55502 55504 55505 55508 55509 55510 55511 55512 From colored 59.111219. —se§d11ngs- colored color- less 0 19 0 25 0 25 0 23 0 25 0 26 0 26 0 25 0 23 0 24 0 5 0 27 0 25 0 21 0 25 o 19 0 25 0 24 0 PA 0 25 0 24 0 25 L79- TABLE XIII-b (continued) From color- lgss kernels -sg§dlings- colored color- less 0 . 24 0 21 0 22 0 21 0 21 0 25 0 20 0 23 0 25 0 20 0 19 0 20 0 20 0 19 0 20 0 15 0 21 0 24 0 20 0 21 0 22 0 23 .1925 (33) Selec- tion No. 55602 55603 55605 55902 55903 55904 55907 55909 56006 56007 56008 56106 56107 56202 56203 56204 56206 56302 56303 56307 56502 56503 'Erom colored kernels From color- lgSB EEEQ§;B -seedlings- -seedling§- colored color— less 0 24 0 25 0 23 0 22 0 25 0 25 0 24 O 25 0 22 0 23 0 21 0 23 0 25 0 24 0 21 0 22 0 23 0 25 0 24 0 27 0 21 0 25 colored color- less 0 24 0 25 0 27 0 21 0 23 O 21 0 19 0 20 0 21 0 23 0 24 0 25 0 21 0 25 0 19 O 20 0 24 0 22 0 23 0 24 0 20 0 20 .1925 (33) Selec- tion NO. 56609 56703 57003 57004 57006 57104 57105 57203 57302 57305 57304 57305 57402 57403 57703 57706 58103 58106 58107 ‘From colored 291219.15 -§gedl;ggs- colored color- less 0 25 0 26 0 25 0 24 0 23 0 25 0 21 0 24 0 25 0 24 0 23 0 24 0 25 0 25 0 25 0 24 0 23 0 25 0 24 0 23 0 22 .TABLE XIII-b (continued) 'From.color- less kergglg -sggd1;ng§- colored color— less 0 20 0 25 0 24 0 25 0 24 0 19 0 15 0 18 0 l9 0 l9 0 20 0 18 0 15 0 l6 '0 l7 0 18 0 20 0 21 0 25 0 21 0 20 ‘1925 .From colored 'From color- (F3) kernels less kernels Selec- -seedliggs- -seed11gg§- tion colored color- colored color- No. less less 58108 0 24 O 25 58303 __g 25 _Q 24 Total ... 0 .. 2504 ..... 0 .. 2261 O -51- TABLE XIII-c. Showing that the colored kernels from 97 3:1 33 ears of the 937§group gave colored and non-colored seedlings in ratio of 2:1 indicating the presence of RSrr and 2625 respective- 1y while the colorless kernels of the same ears gave only colored seedlings indicating the presence of rrrr. 1925 Fran colored kernels From colorless kernels Séfgition colored colorless colored colorless No. seedlings seedlings seedlings seedlings 51601 19 12 5 0 51602 16 7 25 0 51603 19 9 24 0 53403 18 10 23 0 53904 14 5 22 0 53905 23 5 22 0 53906 18 8 22 0 53907 17 9 24 0 53909 21 . 10 17 0 53913 14 7 25 _ 0 53914 16 7 12 0 53916 20 7 17 0 53917 15 9 10 0 53919 15 7 24 0 54005 19 10 24 0 54010 16 6 12 0 54103 18 10 16 0 54104 15 8 18 0 54105 13 5 18 0 54109 20 13 23 0 54111 17 12 11 o "UR- EABLE XIII-c (continued) ‘ 132? Brom colored kernels From colorless kernels Selegtion colored colorless colored colorless No. seedlings seedlings seedlings seedlings 54113 12 7 - 17 0 54114 10 8 20 0 54115 25 14 22 0 54203 16 12 15 0 54204 15 6 24 0 54205 21 16 19 0 54207 13 8 20 0 54208 13 8 19 0 54213 11 6 20 0 54307 11 6 23 0 54309 22 9 20 0 54311 14 12 22 0 54315 14 s 24 ' 0 54406 15 10 22 0 54407 16 9 22 0 54503 13 9 22 0 54506 17 8 23 0 54507 19 7 25 0 54508 18 7 22 0 54510 15 8 25 0 54511 18 8 23 0 54604 15 7 21 0 54606 19 8 23 O v n n i — o- u. .— o—. — an. —- —- —— o.— _ -. —-. n I. _ u ..._ r. — ' . . s 4.. *5- b . ." u s - v I l s 4 - s ..e h. ‘7 . ~. ~ 'a. ‘.A . -. 6. 1 ‘— " i o- . . . 0.- u , s ,‘ .L XVI-n14- -83.. )TABLE XIII-c (continued) %:?? From colored kernels From colorless kernels Selgction colored colorless colored colorless No. seedlings seedlings seedlings seedlings 54704 21 9 25 0 54706 17 8 i 23 0 54707 18 8 24 0 54907 21 9 22 0 54909 18 7 23 0 54910 19 8 25 0 54911 ' 21 8 .21 0 54912 10 4 22 0 55004 17 10 . 14 0 55008 15 9 19 0 55010 18 10 17 0 55404 18 8 25 0 55406 16 10 20 0 55407 15 8 27 0 55412 19 11 24 0 55503 14 6 22 0 55506 20 8 21 0 55507 20 7 24 0 55604 20 11 23 0 55905 21 10 24 0 55906 20 12 23 ', 0 55908 23 10 24 0 55910 19 11 24 O -84.. EABLE XIII-c (continued) -1925 ~From colored kernels 'From colorless kernels Se1jgiion colored colorless colored colorless No. seedlings seedlings seedlings seedlings 56004 16 8 25 0 56005 18 8 24 0 56009 11 5 25 0 56010 23 9 24 0 56102 17 8 21 0 56103 13 10 22 0 56104 14 9 23 0 56105 11 6 21 0 56205 13 9 25 0 56304 23 9 25 0 56305 15 9 26 0 56306 14 8 20 0 56504 14 9 24 ' 0 56505 21 10 22 0 56604 14 9 23 0 56605 18 10 21 0 56606 14 9 21 0 56607 17 10 19 0 56704 19 9 20 0 57005 16 9 21 0 57103 20 10 23 0 57202 27 10 25 0 57305 12 7 21 0 dABLs XIII-c (continued) ‘1925 From colored kernels From colorless kernels ' Se1§glion colored colorless colored colorless No. seedlings seedlings seedlings seedlings 57404 52 17 ' 24 0 57704 23 10 24 0 57705 20 11 _ 25 0 58104 23 12 19 0 58105 17 9 21 0 58302 23 12 25 0 58304 .21 .__1.1 _2_1. .9. TOTAL (97 ears) 11013121.. 16840.. 88100000020830.0000 Expected . 1710 . . . 855 Dev1at10n e “'26 e e 0 +26 .22!;.=._§§_ = 1,5 PoEo 15.1 TABLE XIII-d. Showing that colored seeds of 43 9:7 F5 ears of the 9:7 F2 group gave 2:1 ratio of colored to non- colored seedlings. 1925 1 25 (F3) Number of seedlings (F5) Number of seedlings Selec- Selection tion NO. Colored Non-colored No. Colored Non-colored 51604 22 6 55613 24 15 51606 26 10 55614 24 11 51608 20 11 55616 21 15 53920 20 11 55617 17 11 53925 24 9 55618 16 11 53926 22 8 56015 23 11 53929 18 12 56016 14 11 53931 11 9 56311 13 7 54013 21 13 56312 13 9 54016 20 11 56315 16 11 54018 17 2 56512 12 7 54126 15 7 56513 16 ‘ 9 54127 20 9 56616 20 11 54128 24 9 56710 22 7 54129 23 11 57011 26 12 54133 23 14 57308 15 10 54516 23 12 57309 21 9 54518 18 11 57710 15 9 54610 23 11 57712 14 8 54520 17 10 57713 ____18 .12. Actual total 831 ...... 458 54624 23 12 Theoretical 54710 21 12 total 85903 sees 429.7 Deviation —2%.3 .... .423.3 54712 22 12 22!: : .3§k§. : 2 4 P.E. 11.5 ‘ -8 7.. on the basis of 2:1. This shows a deviation of 28.3 which is 2.4 times its probable error. Obviously, observations coincide with eXpectation fairly well. The results are seen in Table XIII-d. F3 seedling behavior in the 27:57 sh group In this group it should be remembered that the F3 ears are of four kinds of Mendelian ratios. They are 1:0, 3:1, 9:7, and 27:37 ratios. Five of 1:0 53 ears were tested for 5923 and they all gave non— colored seedlings. This is Just what would be eXpected, from results in the group of 1:0 Fa ears from 9:7 E3 ears, and gives substantial support to the hypothesis formulated in diagrams a and b. The results are listed in Table XIVAa. Table XIV;b shows that a total of 117 F3 3:1 ears gave all non- colored seedlings from.both the colored and tie colorless seeds. There- fore, they all carried Rglg (see diagram 0) as expected. It should be recalled that some of these ears had colorless seeds which preved to be the A-tester types and that some of them had colorless seeds which proved to be the C-tester types, as was shown in Table IX-b. Here is another critical test in support of the hypothesis. It was the 2655 factor which caused the C-tester type to be indistinguishable from.the .L—tester type in the seedling tests. The C-tester type would be easily rogued out by the seedling test, if either Rrhr or hgkr were present. Fifty—six 3:1 F3 ears, as shown in Table XIVec, were known as the R-tester types. The colored seeds gave 1051 colored seedlings -88.. $4552 XIV-a. Showing that only non—colored seed- lings resulted from 5 1:0 F3 care which were progenios of the 27:37 F3 group, conclusively indicating the presence of R 23 (110011858) . 1925 (33) . Number of seedlings Selection No. Colored non-colored 51901 0 18 51902 0 19 53801 0 16 57201 0 16 59101 Q 12 T0193]. 0 O O O O O O O O O O O O O O O O 88 -89— TABLE XIV-b. Showing tin-1t 117 F3 ears, progenies of the 27:37 F2 group, gave all non-colored seedlings from both colored and colorless seeds, thereby indicating presence of 2523. 1925 From colored From color- 1925 From colored Fran color- (F5) kernels less kernels (EB) kerngls less mernglg Selec- -s . Mi; Selec- ~seedling;s— m tion colored color- colored color- tion colored color- colored color— No. less less No. ‘ less less 51201 0 25 0 24 52201 0 23 0 26 51202 0 26 0 25 52202 0 22 0 24 51204 0 25 0 26 52204 0 23 0 25 51205 0 26 O 27 52205 0 24 0 21 51304 0 24 0 19 52301 0 22 0 23 51305 0 25 0 20 52401 O 25 0 24 51402 O 23 0 21 52504 0 24 0 25 51403 0 22 0 19 52508 0 21 0 21 51405 0 25 0 20 52602 0 25 0 20 51406 0 24 O 16 52603 0 25 0 21 51502 ‘10 25 O 15 52701 0 26 O 24 51504 0 26 0 6 52702 0 24 0 20 51701 0 25 0 20 52703 0 25 0 21 51703 0 26 0 21 52803 0 20 0 19 51704 0 24 0 22 52902 0 25 0 21 51705 0 22 0 19 53002 0 26 O 23 51706 0 21 O 18 53004 O 24 0 24 51801 0 20 0 20 53005 0 25 0 25 51802 0 25 0 20 53006 0 25 0 20 52002 0 24 0 21 53202 O 25 0 20 52003 0 24 0 20 53203 21 21 52101 0 25 0 25 53205 0 34 0 Z4 . 1925 (Pg) Selec- tion NO. 53304 53306 53501 53601 54801 54802 54805 55103 55202 55203 55205 55302 55303 55304 55305 55307 55702 .Fer colored .From color- .1925 kernels less kernels (F6) -seedllggs- -seedlings- Selec- colored color- colored color- tion less less N0. 0 23 0 23 55703 0 24 O 24 55704 O 25 0 25 55802 0 24 0 21 55803 0 24 0 22 55804 O 25 0 19 56401 0 22 .0 18 56402 0 23 0 19 56403 0 24 0 20 56801 0 25 0 21 56802 0 25 0 23 56804 0 26 O 24 56902 0 27 0 21 56903 0 25 0 25 56905 0 24 0 21 57501 0 22 0 24 57602 0 23 0 25 57603 0 24 0 20 57802 .0 25 o 21 57901 0 26 o 23 58002. 0 24 o 20 58004 0 34 0 19 58201 0 22 0 15 58202 -90- TABLE XIV—b (continued) iron colored kernels From co lor- 1ess kernels -seed1lggs- -§eed1lgg§- colored color- less 0 21 0 22 0 23 O 24 0 23 0 24 0 24 0 25 0 25 0 25 0 24 0 ~26 O 23 0 24 0 25 0 21 0 23 0 24 0 21 0 24 0 25 0 25 0 20 colored color- less 0 17 0 18 0 11 0 19 0 20 0 24 0 24 0 20 0 21 0 23 0 20 0 23 0 24 0 20 0 21 0 l9 0 18 0 15 0 l2 0 l7 0 18 0 l6 0 19 r..- I A _ . .— _. .. .. -. _. — , ‘ . ., . . ‘ l e , . 1 ,5 - ;.‘ . , n ' . u - ... . 0 - ‘u. 1 o 77 < . ._. . » I x . - 7 < o y , . . 1 « . . .,. A 1 s u '1925 (23) Selec- tion NO. 58403 58404 58405 58406 58407 58409 58501 58502 58602 58603 58604_ 58701 58702 58704 58802 58803 58902 58903 58904 'Erom colored -seedliege- -91— I maBLE.XIV;b (continued) 'From color- .1925 \From colored From.color- less 5ezeele (E3) garnels less gezeele -seedliegs- Selec— —seedliegs- -seed;1egs- colored color- colored color- tion Colored color- colored color- less less No. less less 24 0 20 59003 0 23 0 23 25 0 19 59102 0 25 0 19 25 0 20 59103 0 21 0 17 25 0 15 59204 0 25 0 18 21 0 16 59205 0 24 0 21 24 0 17 59206 ___0_ __g_5_ ___g ___ge 23 0 12 Total... 0 ... 2792.... 0 .. 2404 24 0 18 25 0 20 25 0 21 20 0 23 22 0 24 27 0 23 25 0 24 25 0 25 21 0 19 24 0 20 25 0 21 24 0 23 21 0 24 21 o 21 59002 ‘5 EABLE XIVLc. Showing that 56 3:1 F3 ears of the 27:37 F’ group gave colored and colorless seedlings in 8 ratio of 2:1 grom.colored seeds thereby indicating presence or Egrr and R325 respectively and only colored seedlings from colorless seeds thereby indicating rrrr. %§?? From colored seeds From colorless seeds Selegtion colored colorless colored colorless No. seedlings seedlings seedlings seedlings 51203 17 10 20 0 51301 18 10 22 0 51302 21 7 24 0 51303 19 9 22 0 51404 14 6 23 0 51407 23 11 32 0 51413 19 10 27 ,. 0 51503 20 29 30 0 51702 8 4 21 0 51707 18 8 19 0 51803 22 10 15 A 0 51904 13 8 11 0 51905 19 13 22 0 51906 15 9 23 0 52001 18 10 26 0 52203 15 8 25 0 52402 15 10 21 0 52502 25 10 17 0 52503 24 9 34 0 52505 12 7 25 0 52507 15 9 27 0 _9 3- r TABLE 2174c.(Continued) %§:? ' From colored seeds ’ From.colorless seeds Selec- colored colorless colored colorless tion seedlings seedlings seedlings seedlings No. 52510 17 8 18 0 52901 17 9 20 0 52902 13 8 8 0 53003 16 8 7 0 53101 22 14 27 0 53204 12 16 29 0 53206 17 9 25 0 53305 14 8 22 0 53502 13 8 6 0 54804 16 9 17 0 55102 16 8 18 0 55104 18 7 20 . 0 55204 16 ll _ l4 0 55306 16 9 l5 0 55705 13 6 20 0 55805 20 8 21 0 56803 20 9 25 0 56904 19 10 21 0 57803 15 7 25 0 56003 15 6 24 0 58005 11 5 19 0 58203 19 8 23 0 -94- QABLE X1740 (continued) 1925 From colored seeds From colorless seeds (3%) colored colorless colored colorless Selection seedlings seedlings seedlings seedlings No. 58402 31 15 ' 24 0 58408 19 11 24 0 58503 19 12 21 0 58504 12 5 23 0 58505 29 11 23 0 58609 29 15 21 0 58703 27 16 24 0 58801 33 20 20 0 58804 33 15 24 0 58905 24 15 22 0 59001 19 11 23 0 59302 35 15 44 0 592.05 ___Z_L_8_ __;9_ ___2_5_ ‘ __e 0 Actual total 1051 . . . . 571 . . . . . 1215 . . . Expected tOtal (281) 108103 0 s 0 540.? Deviation -30.3 . . . +30.3 2911. = 2919. = 2.3 PoE. 12.7 ..;.: H..- -95.. and 571 non-colored seedlings, where 1081.3:540.7 of colored to non- colored ones were expected. The deviation of 30.3 is 2.3 times its probable error; thus, the observation is fairly close to eXpectancy. The colored seedlings were, naturally, AACGBSr5, and the non-colored, 14883525. All colorless seeds from these ears produced only colored seedlings as rrrr'was present. Their genotype was, of course, AACCrrrr (see diagram b). As shown in Tables XIV;d and 319:8, of 56 9:7 ears 43 gave 835:443 or colored to non-colored seedlings where 852:426 are espected on.a ratio of 2:1 (%E%t-: 1.5), Just as in the case of F2. The remain- ing 13 9:7 F3 ears gave only non-colored seedlings from both colored and colorless seeds. This gives a substantial verification to the hypothesis designated in diagram b (page 70). Theoretically, there are two sorts of 9:7 F3 ears, one with Rgrr, and the other‘mith 2325 in the prOportion of 2:1, hence the 43 9:7 Fs's should be of the fonner type and the remaining 13 should be of the latter type. They are practically in a ratio of 2:1 when the deviation of 5.7 is 2.5 times its probable error (2.3). The conclusion is, then, that observations agree fairlwae 1 with exPectation. Twenty-nine 27:37 F3 ears behaved in the same manner as in F2 giving 652:344 of colored to non—colored seedlings there 664x332 are expected. This results in a deviation of 12, which is only 1.2 times its probable error. Observations are conclusively close to expectancy. The data are arranged in Table XIVLf. Summarizing the above investigation, all data have accorded .-- -9 6.. TABLE XIV;d. Showing that 43 9:7 F3 ears of the 27:37 F group, produced colored and colorless seedlings in the rat 0 of 2:1, thereby indicating the presence of RErr. 1925 1925 (E3) Number of seedlings (EB) Number of seedlings Selec- “ Selec- tion colored non-colored tion colored non-colored N0. NO. 51209 55 22 58505 ’ 17 10 51210 34 18 58618 9 , 4 51212 41 19 58622 13 7 51213 40 21 58625 10 6 51306 23 8 58705 14 10 51308 27 12 58706 8 7 51314 35 14 50711 15 7 51408 35 19 58805 13 6 51409 22 6 58808 16 10 51415 20 9 58809 20 9 51512 12 6 58810 17 10 53806 15 11 58812 16 9 53811 16 9 58813 19 11 53812 19 10 58815 16 9 55310 21 7 58816 12 7 55312 20 10 58817 16 10 56908 14 9 58907 19 10 57205 14 7 58908 14 10 ‘3 57207 23 13 59005 12 11 59009 10 4 57507 22 10 Total (43 ears) ‘Actual 835 443 57509 23 14 Expected 852 426 Deviation -17 ~41? 58412 17 11 92%, : 12 3 1.5 P.h. 11.3 58417 21 11 -97- TABLE XIV;e. Showing that 13 9:7 F ears of the 27:37 F2 group produced only non-colored seedlings from both colored and colorless seeds, thereby indicating the presence 0 f Eglig 0 %§?? From colored seeds From non-colored seeds Selgction colored non-colored colored non-rolored No. seedlings seedlings seedlings seedlings 51207 0 4O 0 19 51208 O 50 0 25 51211 0 62 0 24 51307 0 51 O 26 51522 O 36 O 33 55311 0 32 0 30 56912 0 30 O 26 57810 0 28 O 24 58506 0 17 0 25 58806 0 25 0 . 30 58811 0 20 O 29 59007 0 l7 0 17 58912 _Q .11 .2 .19. Total- 13 ears 0 425 0 , 759 -98- W13 XIV-f. Showing that the colored seeds of 29 27:37 F3 ears of the 27:37 E2 group give colored and non- colored seedlings in a ratio of 2:1. 1925 1925 (F3) Number Of seedlings (36) Number of seedlings Selection Selection 0 NO. colored non-colored No. colored non-colored 51214 38 24 58610 20 5 51217 36 20 58620 12 7 51218 40 17 58630 14 8 51219 38 21 58715 20 9 51220 42 18 58911 7 3 51221 39 23 58913 22 13 51315 23 11 58916 12 8 51316 24 14 58920 13 8 51421 20 12 Actual total 652 . . . 344 Theoretical 51520 15 10 total (2:1) 664 . . . 332 53813 22 10 Deviation -12 . .' . +12 53814 20 9 Dev. : 12 = 1 2 13.2. 10.0 ‘ 55316 22 12 55317 15 17 56814 17 10 56914 17 8 56919 23 10 57213 17 9 57815 17 8 58318 21 ' 9 58415 26 ll -99... with the hypothesis as indicated in diagrams a and b. Therefore, Emerson's A-tester, the male Pl parent in the original crosses, contained figig and its constitution was aaoosgng, and the maternal P1 strains of corn contained rrrr and their constitutions may be ., r r we r r r r written as: AAodr r , Athr r , or AAccr r . SUELAAY l. The work was started in 1923 by the late Professor F..A. Spragg and his associates with an aim to develop a commercial corn, called WA-tester Yellow Dent", for the Michigan firmer. Strains of such corn have been develOped by this present work. 2. In order to develop this type of corn, 24 strains, which were used for corn breeding work here, were crossed with Emerson's ‘A-tester corn which is a white flint having genetic constitution aaCGRR. It was hOped that, in the third generatiOn (33), strains of this new corn, having the genetic constitution saCCRR combined with the yellow dent character from the maternal P1 strains, would be obtained by genetioal breeding processes. 3. .All maternal P1 strains that have been tested sere found to be AA and rr, but were inconsistent for the 0 factor. These results coincide with Dr. Emerson's. Some of the strains were CG, some were Co, and some others had so individuals. Ebr instance, strains of m. A. 0. had all three forms. Their compositions can be written as: AACCrr, AACcrr and AAccrr. Nelson K Bailey and Bailey were found to be AAccrr. 4. 5. 6. -100- bhen these 3 types of corn, AACUrr, AaCcrr and aaccrr were crOSSed.vith.A-tester (aaCGRR), all Fl ears were colored, but they bred differently in Fz'when self—pollinated. The F1 genotype AaCChr from AACCrr X aaC Rh gave ears in 9:7 ratio of colored to colorless seeds, while the Fl genotype haCcRr from AAccrr X aaCCRR produced ears in 27:37 ratios. The cross of AACcrr X.aaCCRR resultdi in 2 Fl genotypes, AaCCRr and AaCcRr. This Kind of F1 ears produced both 9:7 ears and 27:37 ears in the F2 generation in equal numbers. Colored seeds from 9:7 ears and from 27:37 ears bred very differently in F3 when self—pollinated. The former group gave ears with 1:0, 3:1. and 9:7 ratios of colored to colorless aleurones, in preportion of 1:434. Of the 3:1 ears, the colorless seeds were either the R-tester types or the aptester types. The latter types were the ones looked for by the eXperiment at hand. The other group (27:37 3'1 ears) resulted in ears with 1:0, 3:1, 9:7 and 27:37 ratios of colored to colorless alsurones, in prOportion 1:6:1238. The colorless Kernels of the 3:1 ears were of 3 sorts. They'were the A-, C-, and R—tester types. Only the.a-tester types were sought by this experiment, because only from them could the A-tester strains be develOped. The A-tester types could be distinguished from the R-tester types by the seedling test as the latter show anthocyanic pigment in seedlings and could be rOgued out. However, ins Aptester types in the 27:37 Eb group could not be separated out from the C-tester types by this method, because both types resulted in the ears whose .s 7. 8. 9. ~101- seedlings were non-colored. As the multiple allelomorphic factor 8323 was found to be involved in the A-tester types and the C-tester types, the only method of separating these 2 types was to cross them with Ererson's A—, C-, and R-testers in the field. The types which proved to be.A-testers, gave colorless ears when crossed with.A-testers and gave all-colored ears when crossed with C-testers and with R—testers. The C-tester types produced colorless ears when crossed with C-testers and resulted in all-colored ears when crossed with.£-tester and R-tester. The R-tester types produced colorless ears when crossed with.R-testers and gave all-colored ears when crossed with A- and C-testers. In such ways the Aptester types have been picked out from the C-tester types. Some statistical data indicate that these A—tester types of corn which lack the.A factor are not inferior in height, ear length, and ear weight when compared.with the Rptester and the C-tester types which have the A.factor. From 371 selfed F; ears which were A—tester types, 36 with the yellow dent character were considered as desirable for the strains of the A-tester yellow dent corn. or these 36 ears, 11 were selected as the most desirable ears and may be used in producing a possible commercial variety called WA-tester Yellow Dent“. If such a variety is successfully developed from these strains, it will benefit the farmers a great deal, as this corn will aid them in detecting contamination, which is a menace to the pedigree- corn growers. 10. —102- The multiple allelomorphic factors R8 and rr which are partially reaponsible for producing anthocyanins in the plant body were found to be involved in the crosses reported in this eXperiment. Here the factor rr is recessive for aleurone color and dominant for plant color; it aids in producing antho— cyanins in the presence of the A factor. The factor Hg is dominant for aleurone color and recessive fbr plant color; when it is in homozygous condition, it will cause the plants to be non-colored or green even in the presence of the A factor. It was assumed that rr was brought into the cross by the maternal Pl strains (AACCrrrr, AAccrrrr or AACcrrrr) and RS by the ‘A-tester (88033333)- This assumption has been conclusively proved to be correct by the seedling color behaviors in the FE and Eb generations. -105- ~BIBLIOGRLPHY— Bragger, T. 1919. Linkage in maize, the C aleurone factor and waxy endOSperm. Amer. Hat. 53:57-61. Burtty-Javy;J} 1914. Maize, its history, cultivation, handling, and uses. X1 + 831 pp. figs. 1—245. London; Longmans, Green & Co. Collins, G. N. 1916. Correlated characters in maize breeding. Jour. Agr. Research 6:435-54 Czartkowski, Adam. 1914. Anthocyanbildung und Aschenbestandteile. Deut. Bot. Gesell. Bea 32:407-410. Darbishire, A. D. 1911. Breeding and the Mendelian discovery. Vi + 282 pp. 34 figs. 6 pls. New York: Cassell & 00. East, E. M. 1912. Inheritance of color in the aleurone cells of maize. Amer. Eat. 46:563-365. East, E. u. and Hayes, H. K. 1911. Inheritance in maize. Conn..£gr. EXP. Sta. 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Genetic relations of ChlorOphyll and Anthocyanin seedling characters in maize. Genetics 9:91-123. nebber, H. J. 1906. Correlation of characters in plant breeding. Amer. Breeders' Assoc. 2:73-83. wheldale, M. 1911. On the formation of anthocyanin. Jour. Genetics 12133-1580 IA: . «\- _._..._ ‘3‘. tttttt - ' - .1» I a . “4a....”