INHERITANCE OF ECONOMICALLY VALUABLE CHARACTERISTICS IN INTERVARIOUS HYBRIDS OF WHEAT IN SOFT WINTER UNDER FOREST STEPPE Текст научной статьи по специальности «Сельское хозяйство, лесное хозяйство, рыбное хозяйство»

INHERITANCE OF ECONOMICALLY VALUABLE CHARACTERISTICS IN INTERVARIOUS HYBRIDS OF WHEAT IN SOFT WINTER UNDER FOREST STEPPE

Shcherbakova Yu.

Researcher

department of technologies of grain crops NSC "Institute of Agriculture of the NAAS

Abstract

The article presents the results of studies of the inheritance of winter wheat by intervarietal Fi hybrids of valuable economic traits, namely resistance to the main diseases of wheat. Against a natural infectious background, 34% of hybrid combinations demonstrated high resistance to brown leaf rust at the level of 8 points out of the total amount of obtained combinations, of which 5 (14%) reciprocal (direct and reverse) crosses and 7 (20%) direct crosses. Resistance score 7 to the leaf rust pathogen had 20% reciprocal and 29% direct hybrid combinations. 80% of hybrid combinations were moderately resistant to septoria (6 points), of which 13 were reciprocal (37%). With regard to direct combinations, 15 hybrids showed such resistance, that is, 43%. 14 combinations (40%) were highly resistant to powdery mildew pathogen (8 points), of which 11% were reciprocal and 29% were direct. Resistance to powdery mildew of 7 points showed 15 combinations (43%).

Keywords: winter wheat, selection of parental pairs, donor varieties with resistance traits, hybrid combinations, heterosis, dominance.

Introduction

The success of heterotic combinations lies in the successful selection of parental pairs. As noted by S. Boroevich, the most difficult task in combination breeding is the combination of quantitative traits, the genetic structure of which is difficult to study [1].

The degree of phenotypic dominance for assessing the source material at the early stages of the breeding process is used in many agricultural crops [2, 3, 4]. Its effectiveness has been proven in the selection of parental components for crosses, as well as in the evaluation of hybrids [5]. The indicator of the degree of dominance indicates only the inheritance of the studied trait in specific combinations of crossing [6].

Hypothetical heterosis (Ht) shows the excess of the manifestation of the trait in the F1 hybrid over the average value of the parental components. True heterosis - "heterobeltiosis" (Hbt) - makes it possible to identify the strongest manifestation of the trait in F1 in comparison with the best parental form and to assess the breeding value of the hybrid, the possibility of its further commercial use and the high probability of trans-gressive variations emerging from the combination [7].

It should be noted that the magnitude of heterosis in first generation wheat hybrids can vary within wide limits. Its level, found in each specific case, does not always allow predicting the appearance of valuable transgressive forms in the splitting generations. Often, in the first generation of hybrids, the occurrence of in-ter-allelic interaction of genes is possible, which is not transmitted to subsequent generations [8]. Therefore, this indicator should be used in conjunction with other criteria, provides greater efficiency of selection.

According to the researchers, heterosis in winter wheat is observed mainly due to overdominance in terms of productivity elements. This often manifests itself on several signs at the same time [9]. Many authors believe that an indicator of the potential for positive transgressions can be high values of the total combining ability (GCA) of parental components and heterosis in F1 hybrids, which increase the efficiency of selection in the second or third generations [10, 11, 12, 13]. F1 hy-

brids after crosses of common wheat varieties were analyzed to determine the degree of phenotypic dominance of economically valuable traits.

Analyzing the data obtained on the most developed and resistant varieties adapted to the conditions of Polesie and Forest-steppe, based on the available source material of the collection nursery of soft winter wheat, the work was carried out towards the crossing of parental components that are optimal in terms of the combination of quantitative and qualitative traits.

Experimental part

Intraspecific hybridization is the main method in winter wheat breeding. Hybridization is a complex process of the formation of new forms based on the development of the genotype in an environment that is constantly changing. In a hybrid organism, the traits and properties received from the parents, in various combinations, develop anew in each generation. Therefore, it is necessary to know how traits and properties are inherited under certain developmental conditions in order to fully predict the final results of hybridization.

With the aim of transferring valuable traits and properties to the offspring, varieties created in different years, in different scientific institutions, which differ in genetic and geographical origin, duration of the growing season, productivity, grain quality, winter hardiness, drought resistance, lodging resistance and disease resistance, were involved in crosses.

Parental forms of different adaptive and productive potential were involved in crosses. The diallna crossing scheme included many different varieties, but it was the parental forms of the varieties Myronivs'ka 808, Kopylivchanka, Torrild, Kraevyd, Olzhana, Pam'iati Hirka, Kesariia Polis'ka, Benefis, Polis'ka 90 that gave the most viable hybrid combinations with promising characteristics.

The aim of the study is to isolate hybrid combinations of soft winter wheat in terms of resistance to causative agents of major diseases in order to attract them to breeding programs.

The research was carried out in 2014-2015 in the department of breeding and seed production of grain crops of the NSC "Institute of Agriculture of the NAAS"; field experiments were laid in the selection

crop rotation. For two years, soy was the predecessor of wheat. The area of the plots in the Fi hybrid nursery depended on the availability of seeds, a row spacing of 15 cm. Material for the study were 9 varieties of soft winter wheat (Triticum aestivum L.), selected from the working collection of the department of breeding and seed production of grain crops, domestic and foreign breeding, and created on they are based on 35 hybrid combinations. The object of research is soft winter wheat. The subject of research is seeds of varieties and lines of soft winter wheat.

Sowing was done manually at a later date. For the 2015 harvest, sowing was carried out on October 22, 2014, seedlings received on October 29, 2014. The year was unfavorable due to a lack of precipitation and hot weather with significant temperature changes. Due to the autumn drought in 2015, "torn" shoots were obtained for the 2016 harvest, and only in late October -early November (after the appearance of precipitation) did the condition of the plants improve. Disease re-

Inheritance and manifestation of heterosis in F1 hybrids

sistance was assessed using an immunological modification scale for establishing the type of immunity and the intensity of damage to wheat plants, in points, visually.

Field and mathematical-statistical methods were used for research. All plants of each hybrid on the plot were used to describe the qualitative and quantitative traits.

Results and its discussion

Against a natural infectious background, 34% of hybrid combinations demonstrated high resistance to leaf rust, a level of 8 points out of the total number of combinations, of which 5 (14%) reciprocal (direct and reverse) crosses and 7 (20%) direct crosses (Table 1).

Resistance score 7 to the leaf rust pathogen had 20% reciprocal and 29% direct hybrid combinations.

Some of the combinations (17%) had a stability of 6 points. It included only reciprocal combinations: Kesariia Polis'ka/Myronivs'ka 808, Myronivs'ka 808/Pam'iati Hirka, Kraevyd/Pam'iati Hirka, Kraevyd/Myronivs'ka 808.

Table 1

of soft winter wheat for resistance to leaf rust, 2015

Hybrid combinations P1 P2 F1 hp Ht Hbt

9 Myronivs'ka 808 x^ Kesariia Polis'ka 5 6 7 3 27 17

$ Kesariia Polis'ka Myronivs'ka 808 6 5 6 1 9 0

9 Myronivs'ka 808 x^ Pam'iati Hirka 5 5 6 0 20 20

9 Pam'iati Hirka x^ Myronivs'ka 808 5 5 6 0 20 20

9 Pam'iati Hirka x^ Kesariia Polis'ka 5 6 7 3 27 17

9 Kesariia Polis'ka x^ Pam'iati Hirka 6 5 7 3 27 17

9 Pam'iati Hirka x^ Torrild 5 7 8 2 33 14

9 Tomld x^ Pam'iati Hirka 7 5 8 2 33 14

9 Pam'iati Hirka x^ Kraevyd 5 5 7 0 40 40

9 Kraevyd x^ Pam'iati Hirka 5 5 6 0 20 20

9 Kraevyd x^ Kopylivchanka 5 6 7 3 27 17

9 Kopylivchanka x^ Kraevyd 6 5 7 3 27 17

9 Kraevyd x^ Torrild 5 7 8 2 33 14

9 Torrild x^ Kraevyd 7 5 7 1 17 0

9 Kraevyd x^ Myronivs'ka 808 5 5 6 0 20 20

9 Myronivs'ka 808 x^ Kraevyd 5 5 6 0 20 20

9 Torrild x^ Kesariia Polis'ka 7 6 8 3 23 14

9 Kesariia Polis'ka x^ Torrild 6 7 8 3 23 14

9 Myronivs'ka 808 x^ Benefis 5 7 7 1 17 0

9 Kesariia Polis'ka x^ Kopylivchanka 6 6 7 0 17 17

9 Kopylivchanka x^ Myronivs'ka 808 6 5 7 3 27 17

9 Kopylivchanka x^ Pam'iati Hirka 6 5 7 3 27 17

9 Kopylivchanka x^ Olzhana 6 6 8 0 33 33

9 Kopylivchanka x^ Benefis 6 7 8 3 23 14

9 Kopylivchanka x^ Polis'ka 90 6 6 7 0 17 17

9 Kopylivchanka x^ Torrild 6 7 8 3 23 14

9 Torrild x^ Olzhana 7 6 8 3 23 14

9 Torrild x^ Benefis 7 7 8 0 14 14

9 Torrild x^ Polis'ka 90 7 6 8 3 23 14

9 Kraevyd x^ Olzhana 5 6 7 3 27 17

9 Kraevyd x^ Kesariia Polis'ka 5 6 8 5 45 33

9 Kraevyd x^ Polis'ka 90 5 6 7 3 27 17

9 Pam'iati Hirka x^ Olzhana 5 6 7 3 27 17

9 Pam'iati Hirka x^ Benefis 5 7 7 1 17 0

9 Pam'iati Hirka x^ Polis'ka 90 5 6 7 3 27 17

Note: P1 - maternal form; P2 - paternal form; F1 - hybrid; hp - is the degree of phenotypic dominance; Ht - hypothetical heterosis; Hbt - true heterosis.

The most valuable in breeding for resistance are reciprocal hybrid combinations with inheritance by the type of overdominance (hp>+1): Pam'iati Hirka/Kesar-iia Polis'ka, Torrild/Pam'iati Hirka, Kraevyd/Kopy-livchanka, Torrild/Kesariia Polis'ka (direct and reverse), Myronivs'ka 808/Kesariia Polis'ka, and direct Kopylivchanka/Benefis, Kopylivchanka/Myronivs 'ka 808, Kopylivchanka/Torrild, Torrild/Olzhana, Tor-rild/Polis'ka 90, Pam'iati Hirka/Polis'ka 90, Pam'iati Hirka/Olzhana, Kraevyd/Olzhana, Kraevyd/Kesariia Polis'ka, Kraevyd/Polis'ka 90.

Positive hypothetical heterosis (Ht) [14] among the studied sample was observed in all hybrid combinations (100%) and was in the range of 9-45%, while positive true heterosis (Hbt) was observed in 31 (89%) combinations and ranged from 14 up to 40%.

The first generation hybrids and parental components differed in resistance to septoria. 80% of hybrid combinations were moderately resistant to septoria (6 points), of which 13 were reciprocal (37%). Regarding

Table 2

Inheritance and manifestation of heterosis in F1 hybrids of soft winter wheat for resistance to leaf septoria, 2015

direct combinations, such resistance was demonstrated by 15 hybrids, that is, 43% (Table 2).

Moderate susceptibility at the level of 5 points had 20% of combinations, of which 14% were reciprocal: Kesariia Polis'ka/Myronivs'ka 808, Pam'iati Hirka/Torrild, Kraevyd/Kopylivchanka, Kopy-livchanka/Kraevyd, Kraevyd/Pam'iati Hirka, Tor-rild/Kraevyd, Kraevyd/Torrild, Torrild/Kesariia Po-lis'ka, and from direct combinations -Kraevyd/Olzhana, Kraevyd/Polis'ka 90, that is, 2%.

Valuable reciprocal hybrid combinations were obtained with inheritance according to the type of over-dominance (hp>+1): Kraevyd/Torrild, Kraevyd/My-ronivs'ka 808 (forward and reverse), and direct crossing of Kraevyd/Kesariia Polis'ka.

Partial positive dominance (hp=1) had 18 (51%) hybrid combinations, of which 10 were reciprocal and 8. Dominance was not observed in 11 hybrid combinations, 3 of which were reciprocal.

Hybrid combinations P1 P2 F1 hp Ht Hbt

$ Myronivs'ka 808 x^ Kesariia Polis'ka 5 5 6 0 20 20

$ Kesariia Polis'ka Myronivs'ka 808 5 5 5 0 0 0

$ Myronivs'ka 808 x^ Pam'iati Hirka 5 6 6 1 9 0

$ Pam'iati Hirka x^ Myronivs'ka 808 6 5 6 1 9 0

$ Pam'iati Hirka x^ Kesariia Polis'ka 6 5 6 1 9 0

$ Kesariia Polis'ka x^ Pam'iati Hirka 5 6 6 1 9 0

$ Pam'iati Hirka x^ Torrild 6 5 5 -1 -9 -17

$ Torrild x^ Pam'iati Hirka 5 6 6 1 9 0

$ Pam'iati Hirka x^ Kraevyd 6 5 6 1 9 0

$ Kraevyd x^ Pam'iati Hirka 5 6 6 1 9 0

$ Kraevyd x^ Kopylivchanka 4 5 5 1 11 0

$ Kopylivchanka x^ Kraevyd 5 4 5 1 11 0

$ Kraevyd x^ Torrild 4 5 6 3 33 20

$ Torrild x^ Kraevyd 5 4 6 3 33 20

$ Kraevyd x^ Myronivs'ka 808 4 5 6 3 33 20

$ Myronivs'ka 808 x^ Kraevyd 5 4 6 3 33 20

$ Torrild x^ Kesariia Polis'ka 5 4 5 1 11 0

$ Kesariia Polis'ka x^ Torrild 5 5 6 0 20 20

$ Myronivs'ka 808 x^ Benefis 5 6 6 1 9 0

$ Kesariia Polis'ka x^ Kopylivchanka 5 5 6 0 20 20

$ Kopylivchanka x^ Myronivs'ka 808 5 5 6 0 20 20

$ Kopylivchanka x^ Pam'iati Hirka 5 6 6 1 9 0

$ Kopylivchanka x^ Olzhana 5 6 6 1 9 0

$ Kopylivchanka x^ Benefis 5 6 6 1 9 0

$ Kopylivchanka x^ Polis'ka 90 5 5 6 0 20 20

$ Kopylivchanka x^ Torrild 5 5 6 0 20 20

$ Torrild x^ Olzhana 5 6 6 1 9 0

$ Torrild x^ Benefis 5 6 6 1 9 0

$ Torrild x^ Polis'ka 90 5 5 6 0 20 20

$ Kraevyd x^ Olzhana 4 6 5 0 0 -17

$ Kraevyd x^ Kesariia Polis'ka 4 5 6 3 33 20

$ Kraevyd x^ Polis'ka 90 4 5 5 1 11 0

$ Pam'iati Hirka x^ Olzhana 6 6 6 0 0 0

$ Pam'iati Hirka x^ Benefis 6 6 6 0 0 0

$ Pam'iati Hirka x^ Polis'ka 90 6 5 6 1 9 0

An intermediate type of inheritance (-0.5< hp <0.5) of resistance to septoria was observed in reciprocal combinations: Myronivs'ka 808/Kesariia Polis'ka, Torrild/Kesariia Polis'ka (direct and reverse), Kraevyd/Pam'iati Hirka, and direct combinations Kesariia Polis'ka/Kopylivchanka, Kopylivchanka/My-ronivs'ka 808, Kopylivchanka/Polis'ka 90, Kopy-livchanka/Torrild, Torrild/Polis'ka 90,

Kraevyd/Olzhana, Pam'iati Hirka/Olzhana, Pam'iati Hirka/Benefis.

Partial negative inheritance (-1<hp<-0.5) was detected in the reciprocal hybrid combination of Pam'iati Hirka/Torrild.

Positive hypothetical heterosis (Ht) was observed in 30 hybrid combinations (86%) and was in the range of 9-33%, one combination had a negative score and 4 combinations had a score of 0. Positive true heterosis (Hbt) was observed in 12 (34%) combinations and was at 20%. Negative true heterosis was observed in two combinations: reciprocal Pam'iati Hirka/Torrild and direct combination of Kraevyd/Olzhana.

Hybrid combinations differed in resistance to powdery mildew within 6-8 points (Table 3). Highly stable (8 points) were 14 combinations (40%), of which reciprocal 11% and direct 29%.

15 combinations (43%) showed resistance to powdery mildew of 7 points. These are such reciprocal combinations as in Pam'iati Hirka/Myronivs'ka 808, Pam'iati Hirka/Kesariia Polis'ka, Kraevyd/Pam'iati Hirka, Kraevyd/Kopylivchanka, Myronivs'ka 808/Kraevyd, Torrild/Kesariia Polis'ka, Kopy-livchanka/Myronivs'ka 808, Kopylivchanka/Pam'iati Hirka, Kopylivchanka/Torrild, Kraevyd/Olzhana, Kraevyd/Kesariia Polis'ka, and direct combinations -Kopylivchanka/Myronivs'ka 808, Kopy-

livchanka/Pam'iati Hirka, Kopylivchanka/Torrild, Kraevyd/Olzhana, Kraevyd/Kesariia Polis'ka, Kraevyd/Polis'ka 90.

Average resistance to powdery mildew (6 points) had 17% of the combinations. These are reciprocal combinations - Myronivs'ka 808/Kesariia Polis'ka, Pam'iati Hirka/Kraevyd, Kopylivchanka/Kraevyd, Kraevyd/Myronivs'ka 808, Kesariia Polis'ka/Torrild.

Overdominance was observed in 17% of hybrid combinations, partial positive dominance was observed in 31% of combinations. Intermediate dominance was found in 49% of hybrid combinations. Partial negative inheritance (-1<hp<0,5), that is, hybrid depression was in one combination: Kesariia Polis'ka/Torrild.

Table 3

Inheritance and manifestation of heterosis in F1

Hybrid combinations Pi P2 Fi hp Ht Hbt

9 Myronivs'ka 808 x^ Kesariia Polis'ka 6 6 6 0 0 0

$ Kesariia Polis'ka Myronivs'ka 808 6 6 6 0 0 0

9 Myronivs'ka 808 x^ Pam'iati Hirka 6 7 8 3 23 14

9 Pam'iati Hirka x^ Myronivs'ka 808 7 6 7 1 8 0

9 Pam'iati Hirka x^ Kesariia Polis'ka 7 6 7 1 8 0

9 Kesariia Polis'ka x^ Pam'iati Hirka 6 7 8 3 23 14

9 Pam'iati Hirka x^ Torrild 7 7 8 0 14 14

9 Torrild x^ Pam'iati Hirka 7 7 8 0 14 14

9 Pam'iati Hirka x^ Kraevyd 7 5 6 0 0 -14

9 Kraevyd x^ Pam'iati Hirka 5 7 7 1 17 0

9 Kraevyd x^ Kopylivchanka 5 7 7 1 17 0

9 Kopylivchanka x^ Kraevyd 7 5 6 0 -14

9 Kraevyd x^ Torrild 5 7 7 1 17 0

9 Torrild x^ Kraevyd 7 5 7 1 17 0

9 Kraevyd x^ Myronivs'ka 808 5 6 6 1 9 0

9 Myronivs'ka 808 x^ Kraevyd 6 5 7 27 17

9 Torrild x^ Kesariia Polis'ka 7 6 7 1 8 0

9 Kesariia Polis'ka x^ Torrild 6 7 6 -1 -8 -14

9 Myronivs'ka 808 x^ Benefis 6 7 8 3 23 14

9 Kesariia Polis'ka x^ Kopylivchanka 6 7 8 3 23 14

9 Kopylivchanka x^ Myronivs'ka 808 7 6 7 1 8 0

9 Kopylivchanka x^ Pam'iati Hirka 7 7 7 0 0 0

9 Kopylivchanka x^ Olzhana 7 7 8 0 14 14

9 Kopylivchanka x^ Benefis 7 7 8 0 14 14

9 Kopylivchanka x^ Polis'ka 90 7 7 8 0 14 14

9 Kopylivchanka x^ Torrild 7 7 7 0 0 0

9 Torrild x^ Olzhana 7 7 8 0 14 14

9 Torrild x^ Benefis 7 7 8 0 14 14

9 Torrild x^ Polis'ka 90 7 7 8 0 14 14

9 Kraevyd x^ Olzhana 5 7 7 1 17 0

9 Kraevyd x^ Kesariia Polis'ka 5 6 7 3 27 17

9 Kraevyd x^ Polis'ka 90 5 7 7 1 17 0

9 Pam'iati Hirka x^ Olzhana 7 7 8 0 14 14

9 Pam'iati Hirka x^ Benefis 7 7 8 0 14 14

9 Pam'iati Hirka x^ Polis'ka 90 7 7 7 0 0 0

Reciprocal hybrid combinations with inheritance according to the type of superdominance: Myronivs'ka 808/Pam'iati Hirka, Kesariia Polis'ka/Pam'iati Hirka, Myronivs'ka 808/Kraevyd, Myronivs'ka 808/Benefis, Kesariia Polis'ka/Kopylivchanka, Kraevyd/Kesariia Polis'ka.

Positive hypothetical heterosis (Ht) was observed in 77% of hybrid combinations, of which reciprocal 37%, direct 40%.

Heterosis over the best parental component (Hbt) for powdery mildew resistance was found in 46% of hybrid combinations: reciprocal combinations 14% and direct ones 32%. The highest rates of true and hypothetical heterosis were observed in reciprocal combinations - Myronivs'ka 808/Pam'iati Hirka, Kesariia Polis'ka/Pam'iati Hirka, Myronivs'ka 808/Kraevyd and in direct combinations Myronivs'ka 808/Benefis, Kesariia Polis'ka/Kopylivchanka, Kraevyd/Kesariia Polis'ka (Ht = 23-27%, Hbt = 14-17%).

Findings

The most promising for creating varieties resistant to leaf rust are hybrid combinations of Pam'iati Hirka/Torrild, Kraevyd/Torrild, Torrild/Kesariia Po-lis'ka, Torrild/Olzhana, Torrild/Polis'ka 90, Kopy-livchanka/Olzhana, Kopylivchanka/Benefis,

Kraevyd/Kesariia Polis'ka. The most resistant to leaf blight were the combinations Kraevyd/Torrild, Kraevyd/Kesariia Polis'ka, Kraevyd/Myronivs'ka 808. High resistance to powdery mildew was shown by the combinations Myronivs'ka 808/Pam'iati Hirka, Kesariia Polis'ka/Pam'iati Hirka, Pam'iati Hirka/Torrild, Kraevyd/Myronivs'ka 808, Myronivs'ka 808/Benefis, Kesariia Polis'ka/Kopylivchanka, Kraevyd/Kesariia Polis'ka. Complex resistance to two diseases was observed in such hybrid combinations as Kraevyd/Tor-rild, Kraevyd/Myronivs'ka 808, Pam'iati Hirka/Tor-rild, up to three diseases - in the combination of Kraevyd/Kesariia Polis'ka. Thus, these hybrid combinations have a high level of theoretical and hypothetical heterosis with a high degree of dominance and are potentially highly transgressive.

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