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17AGRICULTURAL SCIENCES
COMBINING ABILITY AND GENETIC CONTROL OF THE NUMBER OF FRUITS PER PLANT OF HIGH-LYCOPENE TOMATO IN THE SYSTEM OF DIALLEL CROSSES
Rudas L.,
Candidate of Agricultural Sciences (Ph.D.), Department Head of Plant Growing
Torbanyuk M.
Research Officer, Department of Plant Growing Cherkassy research station of the National Scientific Center «Institute of Agriculture of the National Academy of Agrarian Sciences of Ukraine», UA
Abstract
We studied five parental forms of tomato (Solanum Lycopersicum L.). Were obtained twenty Fi hybrids by the complete diallel scheme (5x5) and used to study inheritance on the trait of the "number of fruits per plant". Were used as parental forms in the system of diallel crosses according to Hayman V.I. and Jinks I.J. : the line №477 (sp, u), variety Alya (sp) and three collection samples with high-lycopene tomato: Dark green (hp -2dg), MO 112 (hp) and T-3627 (Bc). The results of the assessment of the general and specific combining ability of the five parental forms of tomato on the trait of the "number of fruits per plant" are presented, established the character of inheritance. In the course of research it was found that the trait "number of fruits per plant" is controlled by the additive-dominant genetic system. Inheritance of a trait occurs by type of overdominance. The dominant role in genetic control of the trait is played by the dominant effects of genes. Dominant genes determined the increase in trait. The best positive significant effects of general combining ability (GCA) had the line №477, samples Dark green and MO 112. They can be recommended for creation of heterosis hybrids and varieties. From hybrid combinations with high values of specific combining ability (SCA): №477 / T-3627, Alya / Dark green, Alya / MO 112, MO 112 / T-3627 it is necessary to make selections for creation of varieties.
Keywords: tomato, diallel analysis, number of fruits per plant
1. INTRODUCTION
Tomato is a culture of extremely great opportuni-ties[1]. Its fruits are used in salads, pickles and marinades. About 50% of the tomato crop is used to make tomato paste, puree, juice, hot sauces, canning powder [2,3].
The presence of malic and citric acids in fruits improves appetite, activates digestive processes, suppresses harmful microflora[4,5]. Tomato seeds are good for the circulatory system, they dilute the blood and reduce the risk of cardiovascular disease [6]. The peel of the vegetable helps to normalize the gastrointestinal tract and is an excellent prevention of constipation [7,8,9].
Fruit processing waste, consisting mainly of seeds, can be processed into oil. Refined high-quality tomato oil is used for salads. Its digestibility reaches 97% [10,11]. Of value for livestock is tomato cake, which contains 38% crude protein, up to 12 crude fat, 25 - nitrogen-free extractives and about 6% fiber [12,13].
The wide variety of uses of tomato fruits contributes to the constant growth of demand for products[14]. Increasing the production of tomatoes primarily depends on the creation and rapid introduction into production of new hybrids and varieties with the desired economic and valuable traits and high productivity.
High-lycopene varieties and hybrids have the highest nutritional value due to their high content of antioxidants, but are characterized by low productivity [15,16,17]. The components of productivity are the number of fruits per plant and the average weight of the fruit.
The study of the level of heterosis effect and the character of inheritance of these basic economic and valuable traits, allows to predict the selection of parental forms for crosses and expands the possibilities for creating new varieties and hybrids with the desired quantitative and qualitative traits. The best assessment of combining ability can be obtained by dialysis analysis [18-20].
Establishing the characteristics of parental forms by the effects of GCA and SCA, the ratio of their variance in hybrids, as well as the type of gene action (additive or non-additive) is important for predicting the manifestation of breeding and genetic features, determining the effectiveness of selections in hybrid populations by quantitative characteristics [21,22].
2. MATERIALS AND METHODS
The purpose of this research is to identify parental forms with high general and specific combining ability and genetic analysis to establish genetic parameters and the character of inheritance on the trait of the number of fruits per plant.
The research was conducted in 2017-2019 in the fields of breeding and seed crop rotation of Cherkassy research station of the National Scientific Center «Institute of Agriculture of the National Academy of Agrarian Sciences of Ukraine», located in the village Kholodnyansky (49011'N at 31052'E) Cherkassy region, Ukraine. For genetic analysis by (Hayman B.I.,1954; Jinks I.L.,1954) were used the sample №477 (sp, u) and the variety Alya (sp) with a reduced period of fruit ripening and three collection samples with high lycopene content in fruits: Dark green (hp2dg), MO 112 (hp) and T-3627(Bc).
The experiments are based on the method of one-factor experiments [23]. Evaluation of the combining ability on the trait of the "number of fruits per plant" was performed according to the first scheme Griffing B.I. with the matrix of crosses and test p2 (direct and reciprocal crosses + parental forms), where p - is the number of parental forms [18]. Genetic analysis was performed by Hayman V.I. and I.J. Jinks [20, 24].
3. RESULTS AND DISCUSSION
The analysis of variance of a one-factor experiment revealed significant differences between hybrids, that is the number of fruits per plant is different. Differences in combining ability (general or specific) can be expected in the studied lines and varieties. The sum of squares by hybrids is due to genotypic differences. Parent varieties and lines differed in the magnitude of the expression of the trait (Tablel).
Table 1.
Analysis of variance of the number of fruits per plant
Years Type of scattering Sum of squares Degree of freedom Middle square F calc. F Tabl.
General 501,4 74
t- Repetitions 0,1 2
0 Options 465,8 24 19,4* 26,27 1,74
2 Residual 35,5 48 0,7
LSD05 1,43
General 1328,4 74
00 Repetitions 12,3 2
0 Options 1129,6 24 47,07* 12,12 1,74
2 Residual 186,4 48 3,9
LSD05 3,3
General 1639,9 74
OS Repetitions 3,9 2
0 Options 1186,6 24 49,44* 5,28 1,74
2 Residual 449,4 48 9,4
LSD05 5,1
Significant at 5% level
The largest number of the number of fruits per plant was the sample Dark green (7.0-14.7 pieces) and the line №477 (6.0-17.2), the smallest - the variety Alya (4.2-10.7). The average values of the number of fruits per plant in hybrids with the participation of samples
Dark green, MO 112 and line №477 are the largest and are 10.5-19.0, 10.0-19.0 and 9.2-21.6, respectively. With the participation of the variety Alya for three years and the sample T-3627 for two years less than the group average for the parental forms (Table2).
Table 2.
The mean value of the number of fruits per plant in the parental samples, varieties (xp) and hybrids (x F x), days
Variety, sample Years of research
2017 2018 2019
P F1 P F1 P F1
№477 6,0 9,2 14,4 18,4 17,2 21,6
Alya 4,2 8,6 9,9 15,9 10,7 18,0
Dark green 7,0 10,5 14,7 19,0 14,0 19,1
MO 112 6,7 10,0 13,4 16,4 11,8 19,0
T-3627 3,0 7,6 12,1 16,4 13,4 19,0
x 5,4 9,2 12,9 17,2 13,4 18,5
LSDo
1,43
3,25
5,10
The difference between the average F1 and the av- The results of the analysis of variance of the
erage of the parental forms for three years of research combining ability (Table3) indicate significant is positive, that is expression of the trait dominates. differences in general and specific combining ability.
Analysis of variance of the combining ability of the number of fruits per plant
Table 3.
Years Type of scattering Sum of squares Degrees of freedom Middle square F calc. F tabl.
Hybrids 465,8 24 19,4* 26,3 1,79
t— GCA 51,1 4 12,7* 51,9 2,61
0 SCA 101,5 10 10,1* 41,2 2,08
<N Reciprocals 2,7 10 0,3 1,1 2,08
Residual 11,8 48 0,2
Hybrids 1119,3 24 46,6* 12,3 1,79
OO GCA 77,0 4 19,3* 15,2 2,61
O SCA 197,6 10 19,8* 15,6 2,08
<N Reciprocals 98,5 10 9,8* 7,8 2,08
Residual 60,9 48 1,3*
Hybrids 1191,9 24 49,7* 5,3 1,79
OS GCA 75,7 4 18,9* 6,1 2,61
O SCA 291,6 10 29,2* 9,4 2,08
<N Reciprocals 30,0 10 3,0 1,0 2,08
Residual 149,7 48 3,1
Significant at 5% level
Studies have shown that of the number of fruits per plant, the best positive reliable effects of general combining ability (GCA) during the two years of research were the line №477 (1.16-2.32) and the sample Dark green (1.29-1.78), for one year of research - sample MO 112 (0.80) (Table4).
Table 4.
Evaluation of the effects of general combining ability (GCA) on the trait of the number of fruits per plant
„ . _Years_
Sample 2017 2018 2019
№477 0,05 1,16* 2,32*
Alya -0,55* -1,40* -1,26*
Dark green 1,29* 1,78* -0,10
MO 112 0,80* -0,67* -0,25
T-3627_-1,58*_-0,87*_-0,72
LSD05 0,28 0,64 1,00
Significant at 5% level
We found significant differences in specific combining ability (SCA). This indicates that some hybrid combinations within the variety differed significantly from its average value, and in the inheritance of the trait involved non-additive effects of genes. To identify lines and varieties with high or low SCA, for each parent form, we calculated the variance for comparison with the overall average value (Table5).
We found that high reliable values of specific combining ability (SCA) for three years of research
were in the variety Alya (from 1.47 to 6.35), for two years of research in sample T-3627 (3.99-5.22), for one year in sample Dark green (1.76) and line №477 (5.71). The best values of SCA for three years of research were in hybrid combinations №477 / T-3627 (from 1.37 to 3.18), Alya / Dark green (from 2.03 to 4.64); for two years: Alya / MO 112 (1.48-1.53), MO 112 / T-3627 (1.16-3.71).
Table 5.
Estimation of variances of general combining ability (GCA) and specific combining ability (SCA) at an estima_tion of samples and varieties on the trait of the number of fruits per plant of 2017-2019_
Variety, sample Years Alya Dark green MO 112 T-3627 2 5 2 gi
№ 477 2017 2018 2019 -0,08 -2,35* 2,06 1,58* 3,06* 0,24 0,35 1,25 2,14* 1,37* 3,18* 2,29* 0,95 5,71* 1,18 -0,03 1,18 4,97
Alya 2017 2018 2019 2,03* 2,39* 4,64* 1,53* 1,48* 1,72 0,40 2,96* -2,38* 1,47* 4,61* 6,25* 0,27 1,79 1,17
Dark green 2017 2018 2019 1,06* 1,07 -0,65 0,10 -0,44* 0,78 1,76* 3,16 3,31 1,63 3,00 -0,40
MO 112 2017 2018 2019 1,16* -2,36* 3,71* 1,05 1,67 3,09 0,61 0,28 -0,35
T-3627 2017 2018 2019 0,67 5,22* 3,99* 2,46 0,59 0,11
Average value 2017 2018 2019 1,18 4,08 3,56
* Significant at 5% level
Note. Ssi - variance of the effect of specific
combining ability; S 2 - variance of the effect of general combining ability.
Comparison of the variance of the effects of general (S 2) and specific (5Si 2) combining ability revealed that in variety Alya, sample Dark green during three years of studies, samples MO 112 and T-3627 during two years of studies, observed the predominance of dominant effects of genes in genetic control of the number of fruits per plant.
Genetic parameters of the trait
Genetic analysis by (Hayman B.I.,1954) checks the homogeneity of the difference between covariance and variance of Wr-Vr, through Student's t test did not reveal epistatic interaction of genes. Was constructed a graph of the dependence of Wr on Vr according to the indicators of variance and covariance (Fig. 1-3). Graphic analysis shows that the inheritance on the traits of the number of fruits per plant involved additive and non-additive effects of genes (the regression coefficients are reliable and do not differ from one), that is, the regression line does not deviate from the unit slope line. This is confirmed by the high significance of the components D and H1, which characterize the variability of the trait, which is due to the additive and dominant effects of genes (Table 6).
Table 6.
of the number of fruits per plant
Parameters Years
2017 2018 2019
D 2,71±0,85* 2,98±2,07* 2,96±1,68*
F -1,96±2,12* -1,77±5,17* -4,05±4,20*
H1 20,34±2,15* 40,44±5,26* 53,90±4,27*
H2 19,80±2,08* 36,98±5,07* 52,09±4,12*
h2 520,05±1,41* 605,53±3,42* 1214,06±2,7*
E 0,25±0,35 1,27±0,84 3,12±0,69
H1/D 7,51 13,58 18,18
VH1/D 2,74 3,69 4,26
H2/4H1 0,24 0,23 0,24
v4dh1 + f
0,78 0,85 0,72
v4dh1 - f
h2/H2 26,27 16,38 23,31
Conditionally dominant (CD) 8,71 16,48 16,80
Conditionally recessive(CR) 2,66 9,78 10,09
Estimates of H1 and H2 are uneven, from which it is possible to conclude that alleles that positively and negatively determine the trait are unevenly distributed between the parental forms. The uneven ratio of genes with positive and negative effects is indicated by the ratio of H2/4H1, which was 0.23-0.24. The parameter
4adh\ + f
characterizes the ratio of the total
yjAdh 1 - f number of dominant genes to the total number of recessive genes in all parental forms. The index varied in the range of 0.72-0.85, which indicates an excess of recessive gene alleles in the studied samples.
The value of the parameter D, which measures the additive variability in the population, was lower compared to H1, which measures the dominant variability
in three years of research. Values H1> H2, which indicates an unequal ratio of positive and negative effects.
The regression line intersects the negative part of the axis Wr, so we can say that we dominate at all loci. This is evidenced by the value of the average degree of dominance, which was 2.74-4.26.
The table7 shows the indicators of the genetic component F, which reflects the relative contribution of additive and dominant effects of genes in the pheno-typic manifestation of the trait in F1 hybrids. Significant positive effects for three years of research were observed in the sample Dark green, in two years - in the line №477 and the sample MO 112, which indicates the predominance of dominant alleles. Recessive alleles predominated in Alya and T-3627 for three years of research.
Table 7.
Estimation of the direction of dominance (F) of the number of fruits per plant of tomato _for each parental forms and their hybrids_
Samples
№477 Alya Dark green M0112
T-3627
Years
2017
2018
2019
-70,66±17,53* 83,38±17,53* -89,28±17,53* 280,29±17,53* 148,91±17,53*
558,41±73,77* -158,18±73,77* 104,32±73,77* 325,14±73,77* -376,50±73,77*
254,24±100,50* 15,13±100,50* 245,41±100,50* 213,88±100,50* 132,41±100,50*
Significant at 5% level
High correlation coefficients between covariance (Wr) and variance (Vr), as well as regression coefficients b 1 between Wr and Vr (Table 8) indicate that the actual regression lines do not differ significantly from the unit slope.
Table 8.
The results of correlation and regression analyzes of the number of fruits per plant
Indicator 2017 2018 2019
Correlation (r) between Wr and Vr 0,99 0,96 0,41
Regression (b 1) between Wr and Vr 1,11 0,95 1,10
Correlation (r) between хр and Wr and Vr -0,38 0,38 0,95
Regression (b2) between хр and Wr and Vr -0,09 0,05 0,34
The correlation coefficient between the mean values of the trait in the parents ( хр ) and the sum (Wr + Vr) in ranged from minus 0.64 to minus 0.97. This indicates a link between the number of fruits per plant and the presence of dominant genes. At high negative correlations, dominant genes determine the increase in trait.
With the participation of the correlation coefficient between the mean values of the trait of parental
forms in the parents ( хр ) and the sum (Wr + Vr) determined the theoretical values Wdom + Vdom and Wrec + Vrec for varieties and samples, which have dominant and recessive alleles. The variety or sample, which theoretically has all the dominant alleles of the studied parental forms will have Wdom + Vdom 8.71 (2017), 16.48
(2018) and 16.80 (2019). The theoretical value of the parental form with the largest number of recessive genes was 108.35 (2017), 102.52 (2018) and 125.47
(2019).
Using regression graphs (Fig. 1-3), we obtained more complete information about the manifestation of dominant and recessive effects.
In 2017, according to the F1 assessment, line №477, the samples MO 112 and Dark green approached the point with maximum dominance. The point with the highest recessiveness was approached by the variety Alya, which had 75 percent recessive al-leles, and the sample T-3627 with 50 percent recessive alleles.
In 2018, the zone with the highest dominance included the sample of MO 112, as well as all other samples, which had 75 percent of dominant alleles.
Figure 1-3. The graph of dependence Wr by Vr. 1 - Sample №477; 2 - Variety Alya; 3 - Sample Dark green;4 - Sample MO 112; 5 - Sample T-3627.
In 2019, the line №477 and the sample Dark green 112 and T-3627 lie in the middle region of the regres-
were located in the zone with the highest dominance. sion line, far from the points of intersection of the pa-
The point of the variety Alya is located to the zone with rabola with the regression line. the greatest recessiveness, the points of samples MO
4. CONCLUSIONS
Genetic analysis to establish that the trait of the "number of fruits per plant" is controlled by the additive-dominant genetic system. The main role in the genetic control of the trait is played by the dominant effects of genes. Inheritance of a trait occurs by type of overdominance. The high negative value of the correlation coefficient indicates the existence of a relationship between the number of fruits per plant and the presence of dominant genes. At high negative correlations, dominant genes determine the increase in trait.
The best positive and significant effects of general combining ability (GCA) during two years of research had the line №477 (1.16-2.32) and the sample Dark green (1.29-1.78), the sample MO 112 (0.80) in one year of research. They can be recommended for the creation of heterosis hybrids and varieties.
From hybrid combinations with high values of specific combining ability (SCA) for three years of research: №477 / T-3627 (from 1.37 to 3.18), Alya / Dark green (from 2.03 to 4.64); for two years: Alya / MO 112 (1.48-1.53), MO 112 / T-3627 (1.16-3.71) it is necessary to make selections for creation of varieties.
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