Identification of Heterosis and Nature of Gene Action in ...(2016); Saied et al. (2017); Emad et al....
Transcript of 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
Zambrut
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.
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
62
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.
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
63
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).
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
64
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).
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
65
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
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
66
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.
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
67
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
*, ** Significant at 5% and 1% levels of probability, respectively
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
Crosses N D C N D C N D C
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**
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
68
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
*, ** Significant at 5% and 1% levels of probability, respectively
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
*, ** Significant at 5% and 1% levels of probability, respectively
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.
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
69
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
*, ** Significant at 5% and 1% levels of probability, respectively
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
Crosses N D C N D C N D C
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
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
70
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
*, ** Significant at 5% and 1% levels of probability, respectively
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.
5. REFERENCES Abd El-Mohsen, A. A., Abd El-Shafi, M. A., Gheith, E. M. S., and Seliman, H. S. 2015. Using different
statistical procedures for evaluating drought tolerance indices of bread wheat genotypes. Adv. Agric. Biol.
3: 19-30.
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
71
Ahmed, F., Saleem K., Latif A., Hakim K., Ayub K., and Akhtar N. 2011.Genetics of yield and related traits in
bread wheat over different planting dates using diallel analysis. Afr. J. Agric. Res., 6: 1564-1572.
Boyer, J. S. 1982. Plant Productivity and Environment. Science. 218: 443-448.
Cochran, W. G., & Cox, G. M. 1957. Experimental designs. Wiley, New York.
El-Hosary, A. A., Gehan A., and El-Deen, N. 2015. Genetic analysis in the F1 and F2 wheat generations of diallel
crosses. Egypt. J. Plant Breed. 19(2): 355 –373.
Emad F. Morgan, Fawzia I. Moursy, Mohammed S. Abbas, Iman M. Sadek, Mostafa, A. K., and Mohamed M.
A. 2018. Heterosis and Combining ability estimates in 8 X 8 half dillel crosses of bread wheat under two
sowing dates. Egypt. J. Plant Breed. 22: 291-310.
Farooq, J., Khaliq I., Kashif M., Ali, Q., and Mahpara, S. 2011. Genetic analysis of relative cell injury
percentage and some yield contributing traits in wheat under normal and heat stress conditions. Chil. J.
Agr. Res., 71: 511-520.
Farooq, J., Khaliq, J., Akbar, M., Mag, I. V. P., and Hussain, M. 2015. Genetic analysis of some grain yield and
its attributes at high temperature stress in wheat (T. aestivum L.). Annals of R.S.C.B. 21 (3): 71-81.
Fischer, R. A., and Maurer, R. 1978. Drought resistance in spring wheat cultivars. I-Grain yield responses.
Australian J. of Agricultural Research. 29: 897-912.
Gomaa, M. A., El-Banna, M. N. M., Gadalla, A. M., Kandil, E. E., and Ibrahim, A. R. H. 2014. Heterosis,
combining ability and drought susceptibility index in some crosses of bread wheat (Triticum aestivum L.)
under water stress conditions. Middle East Journal of Agriculture Research. 3(2): 338-345.
Griffing, B. 1956. Concept of general and specific combining ability in relation to diallel crossing systems.
Austrlian Journal of Biological Sciences. 9: 463-493.
Gull, S., Aziz, M. K., Ahmed, R. I., Liaqat, S., Rafiq, M., Hussain, F., Rafiq, M. R., and Manjoor, S. A. 2015.
Estimation of heterosis and heterobeltiosis in wheat (T. aestivum L.). Crosses. Basic Res. J. Agric. Sci. and
Review. 4: 151-157.
Jyoti, Yadav. 2017. Genetic architecture of yield and its associated traits in bread wheat (Triticum aestivum L.
em. Thell.) over environments. PhD Thesisin Plant Breeding and Genetics, Department of Plant Breeding
and Genetics, S.K.N. Agriculture University, Jobner. Jaipur.
Kandil, A. A., Sharief, A. E., and Hasnaa, S. M. 2016. Estimation of general and specific combining ability in
bread wheat (T. aestivum L.). Int. J. Agri. R, P37.
Khaled, A. G. A., Motawea, M. H., and Said A. A. 2015. Identification of ISSR and RAPD markers linked to
yield traits in bread wheat under normal and drought conditions. Journal of Genetic Engineering and
Biotechnology. 13: 243–252.
Khan, N., and Naqvi, F. 2011. Effect of Water Stress in Bread Wheat Hexaploids. Curr Rese J of Biol Sci 3:
487- 498.
Kohan, M. Z., and Heidari, B. 2014. Diallel cross study for estimating genetic components underlying wheat
grain yield. J. Biol. Environ. Sci., 8(22): 37-51.
Kumar, D., and Kerkhi, S. A. 2015.Combining ability analysis for yield and some quality traits in spring wheat
(T. aestivum L.). Electronic J. of Plant Breeding. 6 (1): 26.
Leopold, A. C. 1990. Coping with desiccation. In: Stress response in plants: adaptation and acclimation
mechanism. pp: 37-56.
Li, P., Chen, J., and Wu, P. 2012. Evaluation of grain yield and three physiological traits in 30 spring wheat
genotypes across three irrigation regimes. Crop Sci. 52: 110–121.
Ljubicic, N., Petrovic, S., Kostic, M., Dimitrijevic, M., Hristov, N., Kondic-Spika, A., and Jevtic R. 2017.
Diallel analysis of some important grain yield traits in bread wheat crossed. Turkish Journal of Field
crops, 22(1):1-7.
Matzinger, D., and Kempthorne, O. 1956. The modified diallel table with partial inbreeding and intraction with
environment. Genetics. 41: 822-833.
Moghadam, A., and Hadizadeh, M. H.2002. Response of corn hybrids and their parental lines to drought using
different stress tolerant indices. Seed and Plant Journal of Agricultural Research. 18: 255-272.
Saied Abd El-Rahman Shrief, Mohamed Abd El-Maboud Abd El-Shafi, and Sawsan Abd El-Bady El-Sadi.
2017. Heterosis, gene action and combining ability of grain yield and its components in six bread wheat
crosses. Bioscience Research. 14: 1204-1215.
Samir, K. A., and Ismail. 2015. Heterosis and combining ability analysis for yield and it components in bread
wheat (Triticum aestivum L.). Int. J. Curr. Microbio. App. Sci., 4 (8): 1-9.
Semcheddinne, N., Guendouz, A., Oulmi, A., and Hafsi, M. 2017. Screening of Wheat (Triticum durum) for
drought tolerance in semiarid conditions. International Journal of Biosciences. 10(6): 166-178.
International Journal of Agriculture & Agribusiness ISSN: 2391-3991, Volume 3 Issue 1, page 61 – 72
Zambrut
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 ............
72
Shehzad, M., Hussain, S. B., Qureshi, M. K., Akbar, M., Javed, M., Imran, H.M., and Manzoor, S.A. 2015.
Diallel cross analysis of plesiomorphic traits in Triticum aestivum L. genotypes. Genet. Mol. Res., 14(4):
13485-13495.
Singh, D. 1979. Diallel analysis for combining ability over environments. Indian Journal of Genetics and Plant
Breeding. 39: 383-386.
Stanisław Grzesiak, Natalia Hordyńska, Piotr Szczyrek, Maciej T., Grzesiak, Angelika Noga, and Magdalena
Szechyńska-Hebda. 2018. Variation among wheat (Triticum easativum L.) genotypes in response to the
drought stress: I–selection approaches. Journal of Plant Interactions. 14 (1): 30–44.
Steel, R. G., and Torrie, J. H. 1980. Principal and Procedures of Statistics. Mc Grow Hill Book Inc., New York,
USA.
Sundeep Kumar, Jyoti Kumari, Ruchi Bansal, Kuri, B. R., Upadhyay, D., Ashutosh Srivastava, Bhakti Rana,
Manoj, K., Yadav, Sengar, R. S., Amit, K., Singh, and Rakesh Singh. 2018. Multi-environmental
evaluation of wheat genotypes for drought tolerance. Indian J. Genet. 78(1): 26-35.
Yuxiu Liu, Brian, C., Bowman, Yin-Gang Hu, Xi Liang, Weidong Zhao, Justin Wheeler, Natalie Klassen,
Harold Bockelman, J., Michael Bonman, and Jianli Chen. 2017. Evaluation of Agronomic Traits and
Drought Tolerance of Winter Wheat Accessions from the USDA-ARS National Small Grains Collection.
Agronomy. 7, 51; doi: 10.3390/agronomy7030051.
Zambrut Journal, Link Access;
https://zambrut.com
https://zambrut.com/heterosis-wheat/
© Copyright International Journal of Zambrut | Zambrut, Inc.