IL I W“NIH!I!!!“HIWIWIHHNHIIUlHiI‘HHHIHHJ C0000 THF'Q“; This is to certify that the thesis entitled INHERITANCE OF MULTI-PISTILLATE FLOWERING HABIT IN GYNOECIOUS PICKLING CUCUMBER (Cucumis sativus L.) presented by Anand Keshav Nandgaonkar has been. accepted towards fulfillment of the requirements for Master of Science degreein Horticulture @2er Major r Date 2/10/81 0-7639 wuwut. Hut): 25¢ per day per item RETURNING LIBRARY MATERIALS: Place in book return to move charge from circulation records INHERITANCE OF MULTI-PISTILLATE FLOWERING HABIT IN GYNOECIOUS PICKLING CUCUMBER (Cucumis sativus E°) BY Anand Keshav Nandgaonkar A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Horticulture 1981 ABSTRACT INHERITANCE OF MULTI-PISTILLATE FLOWERING HABIT IN GYNOECIOUS PICKLING CUCUMBER (Cucumis sativus E.) BY Anand Keshav Nandgaonkar Progenies of crosses between two multi-pistillate (MP) and two single-pistillate (SP) gynoecious pickling cucumber (Cucumis sativus E.) cultivars were used to de- termine the inheritance of MP expression. From the study it appears that MP is recessive to SP expression. Genetic analysis suggests that one or two major genes with several modifying factors affect this character. The gene symbol proposed for multi-pistillate expression is mp. The genetic information on MP expression should be helpful in cucumber breeding programs. Guidance Committee: This thesis is condensed into a format suited and intended for publication in the Journal of the American Society for Horticultural Science. ii To the memory of my uncle DADA A. N. Kotibhaskar iii ACKNOWLEDGEMENTS The author extends his sincere appreciation to his major professor Dr. J. F. Kelly, for his guidance, encouragement, valuable suggestions and constructive criticism in this manuscript preparation. The author is highly grateful to Dr. L. R. Baker, who served as major professor prior to his leaving the university, for suggesting the problem, inspiring guidance, and valuable suggestions from the beginning of the present investigation til the final shaping of the thesis in its present form. Appreciation is extended to Dr. S. Honma and Dr. K. T. Payne, who extended guidance and appraisal in the manuscript preparation. Special thanks go to Mr. Gene Lester, Michael Dessert, Neil Cowen, James Parrot and Ms. Mary Hunsperger and Nancy Glandon for their invaluable help and active cooperation. He extends his indebtedness and sincere thanks to the Investment in Man, Poona, India for their kind and unforgettable help. iv TABLE OF CONTENTS Page LIST OF TABLES. . . . . . . . . . . . . . . . . . . Vi LIST OF FIGUES O I O C O O O C O O O O Q C C O C . Vii INTRODUCTION. . . . . . . . . . . . . . . . . . . . 1 MATERIALS AND METHODS . . . . . . . . . . . . . . . 5 RESULTS AND DISCUSSION. . . . . . . . . . . . . . . 8 CONCLUSION. . . . . . . . . . . . . . . . . . . . . 24 LITERATURE CITED. . . . . . . . . . . . . . . . . . 25 Table LIST OF'TABLES Characterization of parents for number of pistillate flowers per node (1979-1980) . . Mean number of pistillate flowers per node in 2 MP x 2 SP crosses of gynoecious pickling cucumbers . . . . . . . . . . . . . . . . . Calculated theoretical means and observed means for pistillate flowering habit in F populations of pickling cucumbers derived from gynoecious crosses of MP x SP lines based on one-factor-pair difference . . . . Segregation for single (SP) and multi- pistillate (MP) flower expression in four crosses of gynoecious pickling cucumbers. Genetic analysis of 29 F2 families resulting from four crosses of gynoecious cucumber for flowering habit in the F3 generation. . . . . Mean and range for flower number per node of F3 families from 4 gynoecious cucumber crosses O O O O O O I O O O O O O O O O O O 0 vi Page 19 20 21 22 Figure 1. LIST OF FIGURES A.--Single-pistillate (SP) flower expres- sion in gynoecious cucumber, and B.--Multi- pistillate (MP) flower expression in gynoecious cucumber . . . . . . . . . . . . . Frequency distributions for number of pistil- late (9) flowers per node of parents, F1,F2, BC P and BC P populations in gynoecious cucumber. . l g . . . . . . . . . . . . . . . Frequency distributions for number of pistil- late (2) flowers per node of parents, F1,F , BC P and BC P2 populations in gynoecious cucumber. . I . . . . . . . . . . . . . . Frequency distributions for number of pistil- late (9) flowers per node of parents, F ,Fz, BClP2 populations in gynoecious cucumber. . . Frequegcy distributions for number of pistil- late (+) flowers per node of parents, F1,F2, BC P and BC P populations in gynoecious l 2 cucumber. . . . . . . . . . . . . . . . . . . vii Page 12 14 16 18 INTRODUCTION The development of pickling cucumber (Cucumis sativus E.) cultivars for once-over mechanical harvest has received considerable efforts from plant breeders. Most of the cucumber acreage in Michigan is mechanically harvested; however, greater yields are desirable with once-over harvest. The combination of highly female expression with concentrated fruit-set is a requirement for high yields in once-over harvest systems. Currently used pickling cucumber hybrids produce from one to two fruits per plant for once-over harvest (1,7,8,12). This low fruit-set is due partially to 'first-fruit' inhibition (2,3,9,10,l7). This yield inhibition might be overcome by breeding varieties which simultaneously develop several fruits at the same node. This approach was first suggested by Tied- jens (16) and more recently by Canthfe (2) and Uzcategui and Baker (17). Uzcategui and Baker reported a signifi- cantly greater number of fruits per plant with multi- pistillate (MP) flowers per node as against single-pistillate l (SP) flower per node (Figure l). The greater number of flowers available for pollination possibly increased the number of fruits per plant by circumventing first-fruit inhibition. The development of pickling cucumber hybrids with multi-pistillate flowering might be suited better to once-over harvest than presently used hybrid cultivars with single-pistillate expression. Multiple bisexual flowers per node has been observed by cucumber breeders in hermaphroditic lines, but single- pistillate is common in monoecious and gynoecious lines. Gynoecious and hermaphroditic lines were used to produce gynoecious F recombinants with multi-pistillate flowering 2 (17). The genetics of clustering flowers (multiple- hermaphroditic flowers per node) is conditioned by a single dominant gene closely linked wity hermaphroditic expression (15), which is controlled by the m_locus (4). The purpose of this study was to determine the genetics of single— and multiple-pistillate flowering in gynoecious cucumber lines. Figure l.A.--Single-pistillate (SP) flower expression in gynoecious cucumber. Figure l.B.--Multi-pistillate (MP) flower expression in gynoecious cucumber. Figure 1.A. Figure 1.8. MATERIALS AND METHODS Parental material. The two MP gynoecious lines, 604 G and 598 G described by Uzcategui and Baker (17) were crossed with the two SP gynoecious lines, GY 141 and 551 F2, ,F and BC populations. to produce reciprocal F 2 1 1 All crosses for experimentation were produced in the greenhouse by controlled pollination using standard methods. Genetic analysis. In the spring of 1980, plants of P1,P2,F1,F , and BC populations from the four crosses were 2 1 grown in the greenhouse on raised benches containing a com- mercial peat—based medium. Fourteen-day-old seedlings were transplanted on the greenhouse benches in a randomized com- plete block design with three replications. Five plants each per replicate of the P1,P F and BC P2 generations, 2' l 1 20 plants per replicate of the F2 generation and 10 plants 1GY 14 released by Dr. W. C. Barnes, Clemson Uni- versity, Clemson, S.C. 2551 F released by Dr. H. M. Munger, Cornell Uni- versity, Ithaca, N.Y. per replicate of the BC generation were grown. The 1P1 plants were spaced 30 cm by 48 cm on the bench. The temperature was maintained at 27°:2° C(day) and 21°:2° C (night). The numbers of pistillate flowers per node were recorded for nodes 6 through 15 on the main stem as pre- viously reported (17). Evaluation of F3¢generation. Two to three shoot cuttings were made from randomly selected F plants for 2 all four crosses and were roOted in a mist chamber. All rooted cuttings were self-pollinated to produce F3 seed. In the late summer of 1980, seven F3 populations from each of the 4 crosses and the parental populations were grown as previously described. The number of pistillate flowers per node on the main stem were recorded as mentioned above. Statistical analysis. For all experiments, means and standard deviations were calculated from individual plant data. The reciprocal F populations were not significantly 1 different (p=.05), hence F data were pooled. The method 1 developed by Powers and Locke (11) was used to estimate the number of gene pairs differentiating the parents. 5 (0.75) + i5 (0.25) = E l 2 2 Where fi'is the mean of dominant parent, P2 is the mean of recessive parent, and F2 is the theoretical mean of F2 popula- tion. The chi-square tests were used to determine the goodness of fit of the observed data to the proposed genetic model (14). RESULTS AND DISCUSSION Parental material. The number of pistillate flowers per node were significantly higher for MP than SP lines (Table 1). Between experiments, the actual numbers of pistillate flowers per node varied more (2.6-6.3) for the MP lines, 604 G and 598 G, than for the SP lines, GY l4 and 551 F (0.9-l.l). Environmental conditions influence flowering of cucumber (4,6,13), and appear to play a more significant role in MP for the number of pistillate flowers per node than for SP expression. Table 1. Characterization of parents for number of pistil- late flowers per node (1979—1980).1 Parental Pistillate flowers/node line no. Winter 79 Spring 80 Summer 80 604 G (MP) 5.2a 4.7a 2.6a 598 G (MP) 6.3a 5.7a 4.3a GY 14 (SP) 1.1b l.1b l.0b 551 F (SP) 1.1b 1.1b 0.9b lMean separation within columns by Duncan's multiple range test at 5% level. Genetic analysis. MP and SP expression from the four crosses in this study suggested one or two major genes with several modifying factors affecting this character, SP being dominant to MP expression (Table 2). More than one pistillate flower per node in F1 and BClP2 populations as compared to the dominant parent probably is due to either heterosis (5), or modifying factors. Vigorous hybrid plants probably respond better to environmental conditions than the parental inbred lines (17). The skewness of the F ,F l 2 Table 2. Mean number of pistillate flowers per node in 2 MP x 2 SP crosses of gynoecious pickling cucumbers. moan. (axes Gamma-Eflanuw’ » tion cross 6046 x GY14 598G x GY14 604G x 551E 5986 x 551F (No‘) i _ 5.0. i __ 3.17. 3? _ 5.13. 55 _ 5.13. Pl (MP) 15 4.7 -_+ 0.7 5.7 i 0.7 4.7 t 0.7 5.7 1‘ 0.7 P2 (SP) 15 1.1 t 0.2 1.1 i 0.2 1.1 i 0.2 1.1 i 0.2 F1 30 1.7 i 0.3 1.9 i" 0.4 1.9 1” 0.2 1.9 i' 0.3 F2 60 2.6 1- 1.2 2.3 a; 1.0 2.1 i 0.7 2.6 1- 0.8 BClPl 30 2.8 i 1.1 3.4 i 1.2 2.9 35 1.2 3.7 1. 1.4 BZE’ 15 133: 0J3 ]”6 +(L4 l.84-1.2 l.5:t0.4 1 2 10 and BClP2 populations (Figures 2-5) in all four crosses also supports the hypothesis of one or two major genes with several modifying factors with SP dominant to MP expression. Further evidence is based on the distribution of the BClPl populations. The populations were analyzed using one-gene and two- gene models. For the one-gene model, classification of segregating populations, the class of 2.7-3.2 pistillate flowers per node was selected to divide SP and MP classes. This class approximated the arithmetic mean of the four parents used in this study. This separation of SP and MP classes is also suggested by the low number of individuals falling in this class for BC1 to P1 (MP recessive parent) populations for all four crosses. Classification into two phenotypes, SP and MP, was followed by appropriate testing. The theoretical F2 means were calculated for a one-factor-pair difference using the formula suggested by Powers and Locke (11). The calculated theoretical and observed means for the F populations were 2 not significantly different (Table 3). 11 Figure 2.--Frequency distributions for number of pistillate ( ) flowers per node of parents, F1,F2,BC P and BC P populations from gynoecious MP by P cucumber cross. Frequency (no. plants) 10.. 12 604 G X GY14 (fit P 2 '1 r—J_L—_J——lfi 20. 10.. 20 d 10... 10.. 10.. nclrl Iclr2 1.4 '2.0 ' . l . l . 2 6 3 2 3 8 T 4-4 T 5-0 ' 5-5 ' 6-2 ' Upper class "In for no. 9 flowers Mr node 13 Figure 3.--Frequency distributions for number of pistillate ( ) flowers per node of parents, 'F1,F2,BC P and BC P populations from gynoecious MP by S cucumber cross. Frequency (no. plants) 14 598 G X GY14 I'2 10. .__r O 20... r-—-7 F1 10.) 20. F2 10 .1 1L.__.‘ 0 l___F—_L_L L . MW 102 Icle __J_' 154 T2.0 2.6 I 3.21 3’8 T4.4 l 5.0 I 5'6 I 6’2' liner class vain for no. 9 flowers per node 15 Figure 4.--Frequency distributions for number of pistillate (2) flowers per node of parents, F1,F2 ,BC P and BC 12P populations from gynoecious MP By SP cucumber cross. Frequency (Implants) 16 604GX551F “—"l' r PI 10. . ___, .__..—‘—4_“——. 10- F1 0 'r——1' F2 20.. ,;_____ lOJ——-h 0 0 .L___..———[ Icle 10_ O l I 1'4 1 2'0 2'6 ' 3'2 1 3'8 ' 4'4 1 5°015°6 ' 6-2' Upper class value for no. 9 flowers per node 17 Figure 5.--Frequency distributions for number of pistillate ( F) flowers per node of parents, 2,BC P and BCl P2 populations from gynoecious MP by SP cucumber cross. . Freenency (no. plants) 10) 18 598 G X551 F 2 P1 10... 20.. 10.. 104 aclrl IClPZ 1'4 ' 2-0 ‘ 2-5 ' 3-21 3-8 ' 4.4 ' 5-0 ‘376 ' 6-2 ' Upper class valne for no. 9 flowers per node 6°8 19 Table 3. Calculated theoretical means and observed means for pistillate flowering habit in F2 populations of pickling cucumbers derived from gynoecious crosses of MP x SP lines based on one-factor- pair difference. _: WM— — m I“: Mean flower no./node Cross/F2 P population Theoretical Observed ‘Vaiue 604 G x GY 14 g 2.0 2.6 0.90>P>0.75 598 G x BY 14 2.3 2.3 P>0.99 604 G X 551 F 2.0 2.1 0.99>P>0.95 598 G x 551 F 2.3 2.6 0.90>P>0.75 Goodness of fit was determined for the segregating populations based on a single gene model with SP dominant to MP expression (Table 4). The expected F2 ratio was 3SP:1MP and the expected BC ratio was lstlMP. The P 1P1 values obtained from the segregating populations ranged from 0.05 to 0.95, suggesting good fit to the proposed model. The exception from the one-gene model was noted for the cross 604 G x 551 F. The F2 population did not fit a 3:1 ratio (0.05>P) and BC segregated 2 MP plants when 1P2 none were expected. These two exceptions suggest that there 20 Table 4. Segregation for single (SP) and multi-pistillate (MP) flower expression in four crosses of gynoecious pickling cucumbers. Class Tbtal . Class Expected Population plants frequenc1es ratio P (SP:MP) value (No.) SP MP cross I 604 G (MP) 15 0 15 A11 MP - GY 14 (SP) 15 15 0 All SP - F 30 30 0 1:0 - F% 60 44 16 3:1 0.90>P>0.75 BCIP1 30 17 13 1:1 0.50>P>0.25 cross II 598 G (MP) 15 0 15 A11 MP - G! 14 (SP) 15 15 0 .All SP - F1 30 30 0 1:0 - F2 60 48 12 3:1 0.50>P>0.25 BClPl 30 17 13 1:1 0.50>P>O.25 BCIP2 15 15 0 1:0 - (axeslfil 604 G (MP) 15 0 15 All MP - 551 F (SP) 15 15 0 A11 SP - 512/ 15 15 0 1:0 - F2 60 52 8 3:1 0.05>P>0.025 BClPl 30 12 18 1:1 0.50>P>O.25 BCle 15 13 2 1:0 - cross IV 598 G (MP) 15 0 15 A11 MP - 551 F (SP) 15 15 0 .All SP - F1 30 30 0 1:0 - F2 60 45 15 3:0 P>0.99 BClPl 30 13 17 1:1 0.50>P>0.25 BCIPZI .. 15 15 ~0 ,1:0. . - E/SeedfrcmreciprocalFl (551Fx604 G) wasobtainedtoolate for in- clusion, but later Observation showed no differences in reciprocals. 21 may be two major genes with several modifying factors affect- ing this character. To test this theory, several factorial models were used. Only one model gave a good fit. This model is based on the assumption that 2 genes are involved, with one being completely dominant and another partially segregating ratio of 3:6:3: 2 1:2:1 and a BClPl ratio of 1:2:1:1:2:1 were expected. The chi-square test for these ratios gave a good fit for the dominant. In this model an F cross 604 G x 551 F (0.75>P>0.50). Evaluation of F3 generation. A total of 29 F 3 families from the four crosses were classified as either single pistillate, segregating or multi-pistillate (Table 5). Table 5. Genetic analysis of 29 F2 families resulting from four crosses of gynoecious cucumber for flowering habit in the F3 generation. Class frequencies/class_,_ NO. Cross Families sp/Segregating Multi-pistillate 604 G x GY 14 7 5 1 598 c x or 14 7 7 0 604 c x 551 F 7 5 1 598 G x 551 F 8 4 4 22 3 Mean and range for flower number per node of F families from 4 gynoecious cucumber crosses. Table 6- Pistillate flowers/ F3 F2 Family node (range) no. Cross 1.3 - 3.9 1.3 - 2.7 .2 .9 1.1 1.2 1.2 .9 1.7 1.8 1 604 G x CY 14 0.9 - 1.2 1.0 - 2.3 1.2 1.8 1.0 - 2.0 1.0 - 2.9 1.6 2.3 - 4.1 3.1 90 12 .- 00 ll 32 ll 35 11... CY 14 598 G x 1.4 1.5 05 22 O7 22 1.0 - 4.2 .3 0.9 - 2.2 0.9 - 2.7 1.9 - 3.5 .3 1.4 604 G x 551 F 2.6 1.6 1.7 1.7 2.7 2.9 - 4.2 1.1 - 3.4 1.2 - 4.7 .7 .0 2.2 598 G x 551 F 1.9 7227 O 444 l 95 l 88 32 5 4 4 3 3 5 . 0 1... 1.8 1.0 - .3 2.0 - 4.4 9 23 The data on individual plants in each family are presented in Table 6. The F3 data support the F that multi-pistillate is recessive to the single-pistillate. 2 data which suggest Progenies of multi-pistillate F plants produced mutli- 2 pistillate expression, while progenies from single-pistillate F plants segregated. The segregation as observed in the F 2 3 generation also supports the theory that there may be one or two genes with several modifying factors affecting the pistillate flowering character in gynoecious cucumber culti- vars. CONCLUSION The multi-pistillate habit for gynoecious cucumber lines in this study appears to be recessive to single- pistillate. The genetic analysis suggests that one or two major genes with several modifying factors are affecting this character. The gene symbol proposed for multi- pistillate expression is mp. The genetic information of MP expression should be helpful in cucumber breeding pro- grams. Earlier work (17) suggested that increased number of flowers per node subsequently increased fruit numbers per plant. Through the combination of backcrossing with progeny testing the MP character might be transferred to the parental lines of gynoecious pickling cucumbers. Sub- sequent development of hybrid cultivars with MP expression should result in high yields for once-over harvest. 24 10. LITERATURE CITED Baker, L. R., J. Rudich, J. W. Scott and J. E. Wilson. 1975. Pickling cucumber breeding research. In: Pickle Research at Michigan State University. 1973-1974. Michigan Agr. Ext. 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Res. 36: 721-746e Uzcategui, N. A. and L. R. Baker. 1979. Effect of multiple-pistillate flowering on yields of gynoecious pickling cucumbers. J. Amer. Soc. Hort. Sci. 104: 148-151. Whitaker, J. W. and G. N. Davis. 1962. Cucurbits, Botany, Cultivation and Utilization. Leonard Hill. Ltd. London.