identification of heterosis and nature of gene action in ...(2016); saied et al. (2017); emad et al....

International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72

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Zambrut.com. Publication date: April 19, 2019.

El-Sherbeny, G. A. R., Khaled, G. A. A. & Haitham, M. A. E. 2019. Identification of Heterosis and Nature

of Gene Action in Bread Wheat ............

61

Identification of Heterosis and

Nature of Gene Action in Bread

Wheat Under Normal and

Drought Stress Conditions

El-Sherbeny, G. A. R.1, Khaled, G. A. A.

2 & Haitham, M. A. Elsayed

3

1Prof. El-Sherbeny, G. A. R.,

2Khaled, G. A. A. &

3Haitham, M. A. Elsayed.

Dept. of Genetics, Fac. Agric., Sohag University

Sohag, Egypt

1. INTRODUCTION

Wheat (Triticum aestivum L.) is one of the most important cereal crop overall the world and

Egypt. Abiotic environmental factors are considered to be the main source of yields reductions (Boyer,

1982). Drought is one of the most common environmental stress that affects growth and plant

development through alterations in metabolism and gene expression (Leopold, 1990). Drought stress

may occur early in the season or terminally at grain filling and development.

Abstract: The effects of drought stress on heterosis and the component of genetic variances were

investigated using half diallel mating design among eight bread wheat cultivars. The results

demonstrated that, the majority of cross combinations were earlier, tallest, and high yielding than

their mid and better parents under each environment and their combined data, indicating the

prevalence of heterotic effects and non-additive gene effects. Non-additive gene action (σ2D) was

found to play the major role in the inheritance of studied traits under each environment and their

combined data. Whereas, the interaction of (σ2D x E) were larger than those of (σ2A x E) for all

studied traits, reflecting that the non-additive genetic effects tended to interact with environments

than additive effects. The parents P2 and P8 were a good general combiners for earliness, while

the parents P5, P6, P7 and P8 were the best general combiner for plant height and grain

yield/plant under the two environments and combined data. The crosses (P1xP2), (P2xP3) and

(P2xP5) were the highest desirable specific combining ability effects for days to 50% flowering,

plant height and grain yield/plant under normal conditions. In addition, the crosses (P4xP7),

(P6xP7) and (P2xP5) were the highest specific combining ability effects under drought stress. The

estimates of narrow sense heritability were lower than those of broad sense heritability for days to

50% flowering, plant height and grain yield/plant under normal, drought and combined data,

respectively. The drought susceptibility index based on grain yield/plant exhibited that, the cross combinations which have parents P1, P2, P3 and P8 were relatively tolerant to drought.

Keywords: Wheat, half diallel analysis, drought stress, heterosis, combining ability, gene action and drought susceptibility index.

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Nowadays, Egypt facing a huge problem with the shortage in the water resources and applicable

land for wheat production (Abd El-Mohsen et al, 2015). Therefore, improvement productivity of wheat

cultivars under drought conditions becomes one of the important objectives in wheat breeding program

in arid and semi-arid regions of Egypt. Most of the Egyptian newly reclaimed lands (West and East of

the Delta and West of the Nile Valley in Upper Egypt) suffer from drought and salinity stresses.

Moghadam and Hadi-Zadeh (2002) found that, drought susceptibility index was more useful for

selecting favourable cultivars under stress and non-stress conditions.

Exploitation of heterosis is considered to be one of the outstanding achievements of wheat

breeding. In this trend (Samir and Ismail, 2015; Saied et al. 2017) assessed four different genotypes of

bread wheat using half-diallel design. They reported that the crosses showing the best mid and better

parents could be recommended to improve the days to 50% flowering, plant height and grain

yield/plant.

Diallel cross technique provides useful information on the genetic identity of genotypes

especially on dominance-recessive relations and some other genetic interaction to determinate the

inheritance of traits among a set of genotypes and to identify superior parents for hybrid. Combining

ability analysis of Griffing (1956) partitioned total genetic variance into the variance of general

combining ability GCA, as a measure of additive gene action and specific combining ability SCA, as a

measure of non-additive gene action. The significant and the important role of GCA and SCA for most

studied traits were studied by Kohan and Heidari, (2014); Kumar and Kerkhi, (2015); Kandil et al.

(2016); Saied et al. (2017); Emad et al. (2018). The predominance of non-additive gene effects in the

inheritance of grain yield/plant was reported by Ahmad et al. (2011). On the other hand, additive gene

action controlled the inheritance of plant height and grain yield/plant (Farook et al, 2011; Shehzad et al,

2015).

Therefore, the objectives of the present investigation were directed to study the performance of

eight different genotypes of bread wheat and their half diallel crosses under normal and drought stress

conditions for days to 50% flowering, plant height and grain yield/plant. Moreover, nature of gene

action controlling the inheritance of the three traits was also studied.

2. MATERIALS AND METHODS

2.1 Genetic materials and experimental design:

The genetic materials used in this study were consisted of eight bread wheat genotypes, Misr-1

(P1), Sids-12 (P2), Sahel-1 (P3), Katela (P4), Sakha-94 (P5), Deibera (P6), Weiber (P7) and Canada-462

(P8), which represent a wide range variability in their several agronomic traits. The present study was

carried out at El-Kawther Experimental Research Farm of Faculty of Agriculture, Sohag University,

Sohag, Egypt during the two successive wheat seasons 2016/2017 and 2017/2018.

In the winter season 2016/2017, eight parental genotypes were planted and crossed according to

half diallel mating design to produce 28 F1 hybrids. In the winter season 2017/2018, seeds of eight

parents and their 28 F1 hybrids were sown under normal and drought environmental conditions in a

randomized complete block design (RCBD) with three replications. Each plot consisted of 3 rows 3 m.

long and 30 cm. wide. Plants were spaced by 10 cm. within row. The soil at the experimental site was

sandy to loamy sand. All recommended cultural practise were applied under normal conditions

(irrigation every 10 days) and drought stress (irrigation every 20 days). Data were recorded on ten

plants/genotype chosen at the middle portion of each plot for days to 50% flowering, plant height and

grain yield/plant.

2.2 Biometrical analysis:

In each environment, data were subjected to the analysis of variance to test the significance of

the differences among the tested genotypes according to Cochran and Cox (1957). Combined data over

the two environments were also subjected to the combined analysis of variance to test the interaction of

genotypes with environments.


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Estimates of heterosis over mid and better parents were determined for each cross as follow:

H (M.P) % = x 100

H (B.P) % = x 100

Where;

F1 is the mean of F1 hybrids.

MP and BP: means of the mid parents and better parent, respectively.

The heterotic values were tested for significance to establish the differences of the F1 hybrid

means from their respective mid and better parents using the least significant difference value (L.S.D.)

at 5% and 1% levels of significances, according to the equations suggested by Steel and Torrie (1985).

General combining ability GCA and specific combining ability SCA variances were partitioned

from total genotypic variance in each environment according to (Griffing, 1956) method 2, model 1. In

addition, the combined analysis over the two environments was calculated to partition the men squares

of genotypes and the interaction of genotypes with environments into sources of variations due to

GCA, SCA, and their interaction with the environments (GCA x E and SCA x E).

With the assumption that there is no epistasis, the genetic components could be obtained from

the estimates of GCA variance (σ2g), SCA variance (σ

2s), GCA x E variance (σ

2g x E) and SCA x E

variance (σ2s x E) according to Matzinger and Kempthorne (1956); Singh, (1979). Estimates of

heritability in broad (h2b.s. %) and narrow sense (h

2n.s. %) were also calculated.

Drought susceptibility index “S” estimated according to Fischer and Maurer (1978) equation as

follows:

DSI= [(1-YD/YW) / (1-YMD/YMT)]

Where;

YD: Is the yield under drought stress.

YW: Is the yield under normal condition.

YMD: Is mean yield for all genotypes under drought.

YMT: Is mean yield for all genotypes under normal condition.

Genotypes with average susceptibility or resistance to drought have an “S” value of 1.0. Values of

less than 1.0. Indicate less susceptibility and greater resistance to drought. While, a value of S=0

indicates maximum possible drought resistance (no effect of drought on yield) Fischer and Maurer

(1978).

3. RESULTS AND DISCUSSION

3.1 Genotypic variations:

The analysis of variance (Table 3) showed highly significant between environments for days to

50% flowering, plant height and grain yield/plant with the overall means of normal conditions higher

than those of drought stress conditions. Mean squares of genotypes were found to be highly significant

for all studied traits under the two environments and their combined data, providing evidence for

presence of large amount of genetic variability, which considered adequate for further biometrical

analysis. Moreover, mean squares due to G x E interaction were highly significant for all studied traits,

revealing that these genotypes were inconsistent from environment to another. These results are in

harmony with those of Samir and Ismail (2015); Saied et al. (2017); Semcheddinne et al. (2017); Jyoti

Yadav, (2017); Sundeep et al. (2018); Emad et al. (2018).


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Table 3: Analysis of variances and mean squares of the eight parents and their F1 hybrids for the

studied traits under normal (N), drought (D) conditions and combined data (C).

S.V D.F

Mean squares

Days to 50% flowering Plant height Grain yield/plant

S C N D C N D C N D C

Environments

(E) -- 1 --- ---

190.78

** --- ---

8174.58*

* --- ---

6055.79*

*

Replication (R) 2 -- 10.26 7.79 --- 444.66 2.29 --- 2.35 3.53 ---

Rep. /Treat. -- 4 --- --- 9.02* --- --- 223.47** --- --- 2.94

Genotypes (G) 35 35 61.07*

*

36.04*

*

73.53*

*

148.49

**

130.10

** 250.49**

122.53

**

42.13*

* 123.09**

G x E -- 35 --- --- 23.58*

* --- --- 28.10** --- --- 41.57**

Error 70 140 3.53 3.06 3.29 11.7 3.35 7.52 3.09 2.63 2.86

*, ** Significant at 5% and 1% levels of probability, respectively

3.2 Performance of parents and their crosses:

The results presented in Table 4 indicated that, the performance of the eight parents and their 28

F1 hybrids were variable. The parental average decrease from 94 to 77 days for days to 50% flowering,

102.4 to 78.33 cm. for plant height and 44.53 to 13.6 gm for grain yield/plant under normal conditions

and drought stress, respectively. Drought stress caused reduction about 3.72%, 13.07% and 39.87% for

days to 50% flowering, plant height and grain yield/plant, respectively. It could be noticed that, the best

parents for earliness were P2 and P8 under normal, drought stress and combined data. While, the tallest

were P7 and P6 under each environment and combined data. For grain yield/plant P2, P3, P5 and P6 were

the best under normal, drought stress and combined data, respectively.

The F1 hybrids average reduced from 84.67 to 83.17 days for day to 50% flowering, 103.62 to

91.46 cm. for plant height and 31.33 to 21.69 gm for grain yield/plant in the normal and drought stress,

respectively. The stress conditions caused about 1.77%, 11.74% and 30.77% reduction in the average

of F1 hybrids for days to 50% flowering, plant height and grain yield/plant, respectively. The cross

combinations (P1xP8), (P3xP5), (P5xP8) and (P7xP8) were the earliest hybrids under each environment

and combined data. The tallest cross was (P6xP7) under normal, drought stress and combined data,

respectively. While, the crosses (P1xP3), (P1xP5), (P1xP6), (P2xP5), (P4xP5), (P4xP6), (P4xP8), (P5xP6),

(P5xP7), (P6xP8) and (P7xP8) were the highest for grain yield/plant under each environment and

combined data.

3.3 Drought susceptibility index “S”:

The estimated values of drought susceptibility index “S” based on grain yield/plant for the eight

parents and their 28 F1 hybrids are shown in Table 4. It could be observed that the parental genotypes

P1, P2, P3 and P8 showed S values less than one, revealing relative drought resistance through drought

escape. While, the crosses (P1xP2), (P2xP3), (P3xP5), (P3xP7) and (P3xP8) were relatively tolerant to

drought stress. These results indicated that the tolerant parents P1, P2, P3 and P8 transmitted their genes

controlling drought tolerance to their hybrids. Consequently, these crosses could be considered

promising populations for isolating useful segregates to be cultivated under drought stress. Similar

results found by Khan and Naqvi, (2011); Li et al. (2012); Khaled et al. (2015); Yuxiu et al. (2017);

Stanisław et al. (2018).


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Table 4: Mean performance of the 8 parents and their F1 hybrids for all studied traits under

normal (N), Drought (D) conditions and combined data (C).

Genotypes Days to 50% flowering Plant height Grain yield/plant

DSI N D C N D C N D C

Misr-1 P1 85** 80** 82.50*

* 99.27 83.47 91.37 25.6 22.27 23.93 0.29

Sids-12 P2 79** 78** 78.50*

* 84.53 82.53 83.53 34.87 24.73** 29.8 0.75

Sahel-1 P3 84.33*

* 84**

84.17*

* 96.27 86.4 91.33 35.17 26.40** 30.78* 0.66

Katela P4 90.67 87 88.83 96.27 78.33 87.3 38.10* 20.17 29.13 1.27

Sakha-94 P5 86.67*

*

84.67*

*

85.67*

* 102 86.73 94.37 42.70** 22.07 32.38** 1.45

Diebera P6 87** 87.33 87.17 102.13 91.07*

*

96.60*

* 44.53** 22.53 33.53** 1.54

Weiber P7 94 84** 89 102.3 89.53*

* 95.92* 37.20* 16.1 26.65 1.48

Canada-462 P8 81** 77** 79** 100.2 82.6 91.4 21 13.6 17.3 0.56

P Mean 85.96 82.75 84.36 97.87 85.08 91.48 34.89 20.98 27.94 ----

Reduction % for

Parents Means 3.72% 13.07% 39.87% ----

P1XP2 77** 77** 77** 92.6 86.4 89.5 24.97 20.33 22.65 0.53

P1XP3 84.33*

* 80**

82.17*

* 99.27 89.27 94.27 35.40** 21.3 28.35 1.42

P1XP4 85** 82.67*

*

83.83*

* 87.87 82 84.93 20.7 14.67 17.68 0.66

P1XP5 87* 85.33*

* 86.17* 97.57 86.63 92.1 39.10** 29.60** 34.35** 0.99

P1XP6 85.67*

*

85.33*

*

85.50*

* 101.13 88.2 94.67 36.47** 24.13* 30.30** 1.25

P1XP7 88.33 84.67*

* 86.50* 97.97 89.8 93.88 30.1 20.8 25.45 0.97

P1XP8 78** 79** 78.50*

* 98.3 89.23 93.77 30.33 16.63 23.48 1.38

P2XP3 83.33*

* 85**

84.17*

* 105.33

96.07*

* 100.7 25.97 21.23 23.6 0.54

P2XP4 86.33*

* 90.67 88.5 100 87.87 93.93 36.93** 19.1 28.02 1.77

P2XP5 87.67* 86.33* 87 97.4 89.07 93.23 37.40** 26.20** 31.80** 1.15

P2XP6 87.33* 87.33 87.33 100.47 82.6 91.53 24.37 15.83 20.1 0.9

P2XP7 87.33* 86* 86.67* 105.2 91.27 98.23 31.13 18.8 24.97 1.25

P2XP8 84** 81.33*

*

82.67*

* 108.3 91.8 100.05 26.2 19.1 22.65 0.76

P3XP4 82.67*

*

81.33*

* 82** 103.4

96.60*

* 100 33.1 24.53* 28.82 0.9

P3XP5 79** 84** 81.50*

* 99.2 83.6 91.4 25.03 21.23 23.13 0.45

P3XP6 85.33*

* 88.67 87 101.27 90.2 95.73 27.17 20.9 24.03 0.68

P3XP7 94 81.33*

* 87.67 103.73 83.43 93.58 23.37 18.27 20.82 0.57

P3XP8 80.67*

*

80.67*

*

80.67*

* 107.53 91.47 99.5 26.7 24.03* 25.37 0.34

P4XP5 83** 81.33*

*

82.17*

* 109

100.13

**

104.57

** 32.57 21.7 27.13 1.12

P4XP6 84.33*

*

85.67*

* 85** 100.97 83.03 92 36.43** 24.37* 30.40** 1.23

P4XP7 92.67 80.67*

* 86.67* 106.57 89.13 97.85 26.6 18.77 22.68 0.83

P4XP8 81** 79** 80** 106.43 92.6 99.52 35.20** 26.13** 30.67** 0.95

P5XP6 84.67*

*

85.33*

* 85**

110.73

**

97.80*

*

104.27

** 38.27** 19.2 28.73 1.89


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P5XP7 86** 84.67*

*

85.33*

* 105.87

101.53

**

103.70

** 38.63** 27.20** 32.92** 1.17

P5XP8 79** 80** 79.50*

* 105.4 93.13 99.27 28.97 23.03 26 0.65

P6XP7 94 84.67*

* 89.33

121.60

**

107.60

**

114.60

** 36.67** 26.47** 31.57** 1.05

P6XP8 85** 83.67*

*

84.33*

*

112.93

**

100.60

**

106.77

** 27.53 19.07 23.3 0.89

P7XP8 78** 77** 77.50*

*

115.40

**

99.73*

*

107.57

** 41.83** 24.57* 33.20** 1.72

F1’s Mean 84.67 83.17 83.92 103.62 91.46 97.54 31.33 21.69 26.51 ----

Reduction % for

Hybrids Means 1.77% 11.74% 30.77% ----

L.S.D 5% 3.06 2.84 2.63 5.57 2.98 4.42 2.86 2.64 2.73 ----

L.S.D 1% 4.06 3.78 3.86 7.39 3.96 5.84 3.8 3.51 3.6 ----

Percentage of reduction due to drought stress (R %):

R% P= (M.Pf – M.Ps/M.Pf) X 100.

R% F1= (M.F1f – M.F1s/M.F1f) x 100.

3.4 Estimates of heterosis:

Estimates of heterosis over mid and better parents for studied traits under each environment and

their combined data are presented in Tables 5 and 6. As for days to 50% flowering, the results of

heterosis over mid parents showed that 9, 6 and 7 out of 28 F1 hybrids were significantly earlier than

mid parents under normal, drought stress and combined data, respectively. However, the F1 hybrids

(P1xP2), (P3xP4), (P4xP5), (P4xP8) and (P7xP8) were the best hybrids under both environments and

combined data. Moreover, 3, 3 and 5 out of 28 F1 hybrids were only earlier than better parent under

normal, drought stress and combined data, respectively. While, the cross combination (P4xP5) was the

best desirable over better parent with the heterotic value of -4.23%, -3.95% and -4.09% under normal,

drought stress and combined data, respectively.

Concerning plant height, the results showed that 14, 18 and 15 out of 28 F1 hybrids were

significantly taller than their mid parents under normal, drought stress and combined data, respectively.

Moreover, the tallest hybrids over better parent were 11, 16 and 13 out of 28 crosses with significant

heterotic values under normal, drought stress and combined data, respectively.

Regarding of grain yield/plant, the results showed that 5, 14 and 10 out of 28 F1 hybrids were

significantly out-yielded their mid parents under normal, drought stress and combined data,

respectively. For the better parent, 2, 10 and 10 were the best crosses for grain yield/plant out of 28 F1

hybrids under normal, drought stress and combined data, respectively. These results indicated that, the

majority of crosses were promising for this studied traits and ensures the important of non-additive

gene action in the inheritance of these traits. These results are in harmony with Samir and Ismail, 2015;

Gul et al. 2015; Saied et al. 2017; Jyoti Yadav, 2017.


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Table 5: Estimates of Heterosis (%) over mid parents of 28 F1 hybrids under normal (N),

Drought (D) conditions and combined data (C). Traits Days to 50% flowering Plant height Grain yield/plant

Crosses N D C N D C N D C

P1XP2 -6.09** -2.53* -4.35** 0.76 4.09** 2.34 -17.41** -13.49** -15.69**

P1XP3 -0.4 -2.44* -1.4 1.53 5.10** 3.19 16.51** -12.47** 3.64**

P1XP4 -3.23* -0.99 -2.15 -10.13** 1.36 -4.93* -35.01** -30.87** -33.36**

P1XP5 1.35 3.63** 2.47* -3.05 1.79 -0.83 14.49** 33.51** 22.00**

P1XP6 -0.38 1.98 0.78 0.43 1.07 0.73 4.01** 7.72** 5.47**

P1XP7 -1.31 3.26* 0.84 -2.79 3.82** 0.25 -4.14** 8.42** 0.63

P1XP8 -6.02** 0.64 -2.79* -1.44 7.46** 2.61 30.17** -7.28** 13.89**

P2XP3 2.03 4.94** 3.48** 16.52** 13.74** 15.18** -25.84** -16.96** -22.09**

P2XP4 1.76 9.90** 5.77** 10.62** 9.25** 9.97** 1.22 -14.92** -4.90**

P2XP5 5.83** 6.14** 5.99** 4.43* 5.25** 4.81* -3.57** 11.97** 2.28*

P2XP6 5.22** 5.64** 5.43** 7.65** -4.84** 1.63 -38.62** -33.01** -36.52**

P2XP7 0.96 6.17** 3.49** 12.62** 6.09** 9.48** -13.61** -7.91** -11.53**

P2XP8 5.00** 4.94** 4.98** 17.25** 11.19** 14.39** -6.21** -0.34 -3.82**

P3XP4 -5.52** -4.88** -5.20** 7.41** 17.28** 11.96** -9.65** 5.35** -3.79**

P3XP5 -7.60** -0.4 -4.03** 0.07 -3.43* -1.56 -35.71** -12.39** -26.76**

P3XP6 -0.39 3.50** 1.55 2.09 1.65 1.88 -31.82** -14.57** -25.27**

P3XP7 5.42** -3.18* 1.25 4.48* -5.16** -0.05 -35.42** -14.02** -27.49**

P3XP8 -2.42 0.21 -1.12 9.46** 8.25** 8.90** -4.93** 20.15** 5.53**

P4XP5 -6.39** -5.24** -5.82** 9.95** 21.33** 15.12** -19.38** 2.75* -11.79**

P4XP6 -5.08** -1.72 -3.41* 1.78 -1.97 0.05 -11.82** 14.16** -2.97*

P4XP7 0.36 -5.65** -2.52 7.34** 6.19** 6.81** -29.35** 3.50** -18.68**

P4XP8 -5.64** -3.66** -4.67** 8.34** 15.08** 11.38** 19.12** 54.75** 32.11**

P5XP6 -2.49 -0.78 -1.64 8.49** 10.01** 9.20** -12.26** -13.90** -12.82**

P5XP7 -4.80** 0.39 -2.29 3.64 15.21** 8.99** -3.30* 42.52** 11.54**

P5XP8 -5.77** -1.04 -3.44** 4.25 9.99** 6.87** -9.04** 29.13** 4.67**

P6XP7 4.42** -1.17 1.41 18.96** 19.16** 19.05** -10.27** 37.04** 4.92**

P6XP8 1.19 1.83 1.49 11.63** 15.85** 13.59** -15.98** 5.56** -8.32**

P7XP8 -10.86** -4.35** -7.74** 13.98** 15.88** 14.85** 43.75** 65.46** 51.08**

LSD 5% 2.21 2.06 2.11 4.03 2.15 3.19 2.06 1.91 1.96

LSD 1% 3.16 2.94 2.98 5.75 3.06 4.52 2.94 2.73 2.77


Table 6: Estimates of Heterosis (%) over better-parent of 28 F1 hybrids under normal (N),

Drought (D) conditions and combined data (C). Traits Days to 50% flowering Plant height Grain yield/plant


P1XP2 -2.53 -1.28 -6.67** -6.72** 3.51** -2.05 -28.39** -17.79** -23.99**

P1XP3 0 0 -0.4 0 3.32** 3.17 0.65 -19.32** -7.89**

P1XP4 0 3.34* 1.61 -11.48** -1.76 -7.05** -45.67** -34.13** -39.31**

P1XP5 2.35 6.67** 4.45** -4.34* -0.12 -2.41 -8.43** 32.91** 6.08**

P1XP6 0.79 6.67** 3.64** -0.98 -3.15* -1.99 -18.10** 7.10** -9.63**

P1XP7 3.92** 5.84** 4.85** -4.23* 0.3 -2.13 -19.09** -6.60** -4.50**

P1XP8 -3.70** 2.59* -0.63 -1.89 6.90** 2.59 18.48** -25.33** -1.88

P2XP3 5.48** 8.97** 7.22** 9.41** 11.19** 10.26** -26.16** -19.58** -23.33**

P2XP4 9.28** 16.24** 12.74** 3.88 6.47** 7.59** -3.07** -22.77** -5.97**

P2XP5 10.98** 10.68** 10.82** -4.51* 2.69* -1.21 -12.41** 5.94** -1.79

P2XP6 10.54** 11.96** 11.25** -1.63 -9.30** -5.25** -45.27** -35.99** -40.05**

P2XP7 10.54** 10.26** 10.41** 2.84 1.94 2.41 -16.32** -23.98** -16.21**

P2XP8 6.33** 5.62** 5.31** 8.08** 11.14** 9.46** -24.86** -22.77** -23.99**

P3XP4 -1.97 -3.18* -2.58* 7.41** 11.81** 9.49** -13.12** -7.08** -6.37**

P3XP5 -6.32** 0 -3.17* -2.75 -3.61** -3.15 -41.38** -19.58** -28.57**

P3XP6 1.19 5.56** 3.36* -0.84 -0.96 -0.9 -38.99** -20.83** -28.33**

P3XP7 11.47** -3.18* 4.16** 1.39 -6.81** -2.44 -37.18** -30.79** -32.36**

P3XP8 -0.41 4.77** 2.11 7.32** 5.87** 8.86** -24.08** -8.98** -17.58**

P4XP5 -4.23** -3.95** -4.09* 6.86** 15.45** 10.81** -23.72** -1.68 -16.21**

P4XP6 -3.07* -1.53 -2.49* -1.14 -8.83** -4.76** -18.19** 8.17** -9.34**

P4XP7 2.21 -3.96** -2.43* 4.17* -0.45 2.01 -30.18** -6.95** -22.14**


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P4XP8 0 2.59* 1.27 6.22** 12.11** 8.88** -7.61** 29.55** 5.29**

P5XP6 -2.31 0.78 -0.78 8.42** 7.39** 7.94** -14.06** -14.78** -14.32**

P5XP7 -0.77 0.79 -0.39 3.49 13.40** 8.11** -9.53** 23.24** 1.67

P5XP8 -2.47 3.89** 0.63 5.19* 7.38** 5.19** -32.16** 4.35** -19.70**

P6XP7 8.05** 0.79 2.48* 18.87** 18.15** 18.63** -17.65** 17.49** -5.85**

P6XP8 4.94** 8.66** 6.75** 12.71** 10.47** 10.53** -38.18** -15.36** -30.51**

P7XP8 -3.70** 0 -1.89 12.81** 11.39** 12.15** 12.45** 52.61** 24.52**

LSD 5% 2.55 2.38 2.44 4.64 2.48 3.69 2.39 2.19 2.27

LSD 1% 3.63 3.39 3.45 6.62 3.54 5.22 3.42 3.13 3.22


3.6 Combining ability analysis:

Combining ability analysis of variance (Table 7) showed that GCA and SCA mean squares for

studied traits were highly significant under each environment and their combined data, confirming the

important role of all types of gene actions in the expression of these traits. While, the ratios of

GCA/SCA were found to be larger than unity for all studied traits except grain yield/plant under each

environment and combined data. In addition, the interaction of GCA x E was only highly significant for

days to 50% flowering. While, the interaction of SCA x E was only highly significant for grain

yield/plant. Furthermore, the ratios GCA x E/ SCA x E were more than one for all studied traits except

grain yield/plant, suggesting that the magnitudes of all types of gene actions fluctuated from normal to

drought stress conditions. These results were in agreement with the results obtained by Gomaa, et al.

(2014); Kumar and Kerkhi (2015); Samir and Ismail. (2015); Jyoti Yadav, (2017); Saied et al. (2017).

Table 7: Combining ability analysis of variance for all the studied traits under normal (N) and

drought (D) conditions as well as their combined data (C).

S.V D.F

Mean squares

Days to 50% flowering Plant height Grain yield/plant

S C N D C N D C N D C

GCA 7 7 68.43** 34.61*

*

81.66*

*

180.18*

*

101.62*

* 267.2 30.51** 5.99 26.7

SCA 28 28 8.34 6.36 10.22*

* 16.83** 28.80** 37.57 43.43** 16.10** 44.61**

GCA x E --- 7 --- --- 64.13*

* --- --- 43.79 --- --- 29.41

SCA x E --- 28 --- --- 13.45 --- --- 24.18 --- --- 44.61**

Error 70 140 1.18 1.02 1.1 3.9 1.12 2.51 1.03 0.88 0.95

GCA/SCA 8.21 5.44 7.99 10.71 3.53 7.11 0.7 0.37 0.59

GCA x E/SCA x E --- --- 4.77 --- --- 1.81 --- --- 0.66


3.7 GCA effects (gi):

The results in Table 8 showed that, the parental genotypes P2 and P8 exhibited negative general

combining ability effects toward earliness under both environments and their combined data. In

addition, the parental genotypes P7 and P8 were the best general combiners for tallness under each

environment and combined data. Regarding of grain yield/plant, the parental genotypes P5 and P6

recorded the best general combiner under each environment and their combined data.

It could be concluded that the parents P5, P6, P7 and P8 were found to be excellent combiners for

the majority of studied trait under the two environments and their combined data. Consequently, these

promising parents could be utilized in wheat breeding program to improve studied traits under each

environment and combined data.


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Table 8: General combining ability effects for all the studied traits under normal (N), drought

(D) conditions as well as their combined data (C). Traits Days to 50% flowering Plant height Grain yield/plant

Genotypes N D C N D C N D C

Misr-1 P1 0.81* -0.57 0.12 -3.89** -4.62 -4.26 1.57** -0.36 0.60**

Sids-12 P2 -2.33** -1.87** -2.09 -6.93 -2.99 -4.96 -1.23** -0.12 -0.67**

Sahel-1 P3 0.34 1.8** 1.07** -0.35 0.003 -0.18 -1.05** -0.46 -0.76**

Katela P4 0.18 1.00** 0.59** -1.84** -2.03** -1.94 -2.49 -0.63* -1.56

Sakha-94 P5 -0.03 0.67 0.02 0.7 0.07 0.39 1.85** 1.8 1.83**

Diebera P6 -0.06 2.20** 1.07** 1.76** 1.50** 1.63** 2.4 0.26 1.33

Weiber P7 5.08 0.67 2.87 5.67 5.01 5.34 -0.12 -0.18 -0.15

Canada-

462 P8 -3.99 -3.3 -3.65 4.88 3.05 3.97 0.95* -0.3 -0.63**

SE(gi) 0.32 0.29 0.16 0.58 0.31 0.23 0.29 0.28 0.14


3.8 SCA effects (Sij):

The results in Table 9 indicated that, the best crosses for days to 50% flowering were (P1xP2),

(P3xP7) and (P6xP7) under normal condition, (P2xP3), (P4xP7) and (P5xP7) under drought stress and

(P1xP2), (P4xP5) and (P6xP7) under combined data. The highest desirable SCA effects toward tallness

were obtained from the crosses (P1xP5) and (P2xP3) under normal condition and combined data,

respectively. Moreover, the cross (P2xP6) was the highest under normal condition, and (P6xP7) and

(P7xP8) were the highest under drought stress. In addition, the cross (P1xP6) was the highest SCA

effects toward tallness under drought stress and combined data. Regarding to grain yield/plant, the

crosses (P1xP3), (P2xP5) and (P6xP7) exhibited desirable SCA effects for increasing grain yield per plant

under drought stress and combined data. Moreover, the cross combination (P1xP6) recorded the best

SCA effect under normal and combined data. In addition, the best SCA effects were (P1xP4), (P1xP5)

and (P3xP4) for normal condition and (P1xP2) for drought stress for the same trait.

Table 9: Specific combining ability effects for all the studied traits under normal (N), drought (D)

conditions as well as their combined data (C). Traits Days to 50% flowering Plant height Grain yield/plant


P1XP2 -4.44** -2.64** -3.54 -6.99** 0.11 -3.45** 2.39** 3.69** 3.04

P1XP3 -1.77 -0.31 -1.04 -1.84 0.98 0.43 2.53** 5.69 4.11

P1XP4 4.73 3.49** 4.11 -0.35 -5.06 -2.70** 6.89 -0.37 3.27

P1XP5 0.93 2.09** 1.51* 2.84 1.25 2.04* 7.16 -0.89 3.13

P1XP6 1.29 2.63** 1.96** 1.92 4.15** 3.03** 8.44 1.1 4.77

P1XP7 3.16** 0.83 1.99** -1.82 -0.89 -1.36 3.63** -4.88 -0.63

P1XP8 -0.77 -2.21* -1.49** -3.14 -5.87 -4.50** -11.74 -7.26 -9.5

P2XP3 1.36 -3.01** -0.82 4.20* 2.21** 3.21** 5.55 0.35 2.95

P2XP4 2.19* 0.46 1.33* -5.71** -3.02** -4.37** -7.71 -6.11 -6.91

P2XP5 4.39** 4.06** 4.23 1.45 -0.49 0.48 6.35 6.39 6.37

P2XP6 3.09** 1.93* 2.51** 3.96* -0.35 1.8 3.16** 2.46** 2.81**

P2XP7 0.63 2.79** 1.71* -3.12 -2.26** -2.69** -0.68 -0.42 -0.55

P2XP8 -0.64 1.09 0.23 -1.99 -0.87 -1.43 0.38 -4.47 -2.05**

P3XP4 0.86 4.79 2.83** -0.15 -0.15 -0.15 8.36 -1.34 3.51

P3XP5 2.39** 1.39 1.89** -5.29** -1.05 -3.17** 4.48 3.33** 3.91

P3XP6 2.09* 0.26 1.18 -3.28 -8.94 -6.11 -9.1 -5.5 -7.3

P3XP7 -3.04** 0.46 -1.29 -2.46 -3.79** -3.12** 0.19 -2.09** -0.95

P3XP8 2.69** -0.24 1.23 1.43 -1.29 0.07 -3.92** -1.67* -2.79**

P4XP5 -6.1 -0.14 -3.12 -2 -4.48 -3.24** -6.45 -1.46 -3.96

P4XP6 0.26 2.39** 1.33* -0.99 0.69 -0.15 -4.87 -0.26 -2.56**

P4XP7 3.79** -3.41** 0.19 -2.43 -9.59 -6.01 -6.15 -2.45** -4.29

P4XP8 -0.47 -0.11 -0.29 2.15 0.39 1.28 -1.98* 3.44** 0.73

P5XP6 -0.54 0.33 0.11 -3.84* -8.58 -6.21 0.06 0.77 0.42

P5XP7 2.66** -3.14** -0.24 -2.15 -5.99 -4.07** -7.25 -4.38 -5.82

P5XP8 0.06 -0.84 -0.39 -1.49 -0.56 -1.03 2.18* 3.11** 2.64**

P6XP7 -3.97** -1.27 -2.62** -3.90* 4.98 0.54 4.23 5.59 4.91


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P6XP8 0.06* -1.97** -1.94** -3.58* -1.46 -2.52** -4.61 1.54* -1.53*

P7XP8 -1.04 3.23** 1.09 0.04 2.49* 1.27 -3.52** -1.98* -2.75**

SE(Sij) 0.98 0.92 0.67 1.79 0.96 1.02 0.92 0.85 0.63


It could be observed that the promising hybrids were resulted from the crossing (good x good),

(good x poor) and (poor x poor) general combiners. Therefore, it is not necessary that parents having

high estimates of GCA effects would also give high estimates of SCA effects in their respective

crosses. In general, the promising crosses which showed desirable SCA effects gave also high estimate

of useful heterosis as previously mentioned. These finding indicate that non-additive gene action

played an important role in the inheritance of these traits. The same results were obtained by Gomaa, et

al. (2014); Kohan and Heidari (2014); Jyoti Yadav, (2017); Saied et al. (2017).

3.9 Estimates of genetic parameters:

The genetic parameters included additive (σ2A) and non-additive (σ

2D) genetic, as well as the

values of heritability in broad sense (h2

b.s %) and narrow sense (h2

n.s %) are presented in Table 10. The

results indicated that, the magnitudes of the additive genetic variance (σ2A) were larger than those of

non-additive ones (σ2D) for days to 50% flowering under each environment and combined data.

However, the magnitude of σ2A x E interaction was less than σ

2D x E for the same trait. The estimates

of broad sense heritability (94.22, 91.51 and 34.58%) were larger than those of narrow sense

heritability (59.03, 47.03 and 33.70%) under normal, drought stress and combined data, respectively. In

addition, the estimates of σ2A for plant height were higher than those of σ

2D under normal condition

and combined data. While, the magnitude of σ2A was lower than σ

2D under drought stress. Moreover,

the magnitude of σ2A x E interaction was less than σ

2D x E for this trait. In addition, the values of

broad sense heritability were 92.12%, 97.43% and 55.80% under normal, drought stress and combined

data, respectively. While, the estimated values of narrow sense heritability for plant height were

66.00%, 33.58% and 48.05% under each environment and combined data, respectively. Concerning to

grain yield/plant, the magnitudes of σ2A were lower than σ

2D under each environment and combined

data Furthermore, the magnitudes of σ2A x E interaction was less than σ

2D x E for this trait. Moreover,

the estimates of broad sense heritability was larger many times than those of narrow sense heritability

under each environment and their combined data. These results are agree with those obtained by Kohan

and Heidari (2014); Farooq et al. (2015); El-Hosary et al. (2015); Kandil et al. (2016); Ljubicic et al.

(2017); Saied et al. (2017).

Table 8: Genetic components for all the studied traits under normal (N), drought (D) conditions

as well as their combined data (C).

Genetic Components Days to 50% flowering Plant height Grain yield/plant

N D C N D C N D C

σ2 A 12.02 5.65 5.45 32.67 14.56 22.31 2.58 2.02 1.29

σ2 D 7.16 5.34 2.87 12.93 27.69 14.76 42.4 15.18 14.87

σ2 A x E -- -- 3.38 -- -- 1.31 -- -- 1.01

σ2 D x E -- -- 3.39 -- -- 5.55 -- -- 13.92

σ2 e 1.18 1.02 1.1 3.9 1.12 2.51 1.03 0.88 0.95

Narrow h2

n.s% 59.03 47.03 33.7 66 33.58 48.05 5.61 11.17 4.27

Broad h2

b.s% 94.22 91.51 34.58 92.12 97.43 55.8 97.76 95.15 51.69

4. CONCLUSION

It could be concluded that, according to estimates of heterosis and nature of gene action, plant

breeder could use days to 50% flowering and grain yield/plant as indicators which could be used for

selection favourable genotypes to cultivate under drought stress conditions.

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