Scholarly article on topic 'Modified model for assessment of maternal effects in first generation of faba bean'

Modified model for assessment of maternal effects in first generation of faba bean Academic research paper on "Agriculture, forestry, and fisheries"

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{"Faba bean (Vicia faba L.)" / "Griffing’s method" / Hayman / "Full-diallel crosses" / GCA / SCA / "Reciprocal effect" / "Maternal effect"}

Abstract of research paper on Agriculture, forestry, and fisheries, author of scientific article — Zeinab E. Ghareeb, W.M. Fares

Abstract This study was carried out during two successive growing seasons, 2013/14 and 2014/15 at Giza Research Station, ARC, Giza, Egypt, in order to assess proposed modified model when the general and specific combining ability (GCA and SCA) effects were portioned to female and male effects and to assess the maternal and reciprocal effects in 5×5 diallel crosses of faba bean. The results of statistical analysis for parental genotypes and their F 1 hybrids, revealed highly significant differences among them for all studied traits except number of branches per plant. The cytoplasmic components were significant for number of seeds/plant, weight of 100 seeds and seed yield/plant. Results also revealed that estimated GCA effects according to Griffing’s method were equal to the average of GCA effects of each parent, after partitioning in the proposed model. In addition, the average of the difference between female and male GCA effects would provide valid and precise estimation of the maternal effect (favorable alleles, which are mainly additive) as previously confirmed by Hayman analysis for number of seeds/plant, weight of 100 seeds and seed yield/plant. The SCA effects calculated according to Griffing’s method equaled the average of SCA effects of each cross and its reciprocal. Meanwhile, in the proposed model, the average of the difference between SCA effects of each cross and its reciprocal equaled the reciprocal effects. This would prove that reciprocal effect provides precise estimation to the interaction effect between nuclear and cytoplasmic genes of the cross and its reciprocal hybrid.

Academic research paper on topic "Modified model for assessment of maternal effects in first generation of faba bean"

Annals of Agricultural Science (2016) 61(1), 77-85

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Faculty of Agriculture, Ain Shams University Annals of Agricultural Science

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Modified model for assessment of maternal effects ck»^ in first generation of faba bean

Zeinab E. Ghareeb *, W.M. Fares

Centeral Laboratory for Design and Statistical Analysis Research, ARC, Giza, Egypt

Received 18 October 2015; accepted 23 January 2016 Available online 1 June 2016

KEYWORDS

Faba bean (Vicia faba L.); Griffing's method; Hayman;

Full-diallel crosses;

Reciprocal effect; Maternal effect

Abstract This study was carried out during two successive growing seasons, 2013/14 and 2014/15 at Giza Research Station, ARC, Giza, Egypt, in order to assess proposed modified model when the general and specific combining ability (GCA and SCA) effects were portioned to female and male effects and to assess the maternal and reciprocal effects in 5 x 5 diallel crosses of faba bean. The results of statistical analysis for parental genotypes and their F1 hybrids, revealed highly significant differences among them for all studied traits except number of branches per plant. The cytoplasmic components were significant for number of seeds/plant, weight of 100 seeds and seed yield/plant. Results also revealed that estimated GCA effects according to Griffing's method were equal to the average of GCA effects of each parent, after partitioning in the proposed model. In addition, the average of the difference between female and male GCA effects would provide valid and precise estimation of the maternal effect (favorable alleles, which are mainly additive) as previously confirmed by Hayman analysis for number of seeds/plant, weight of 100 seeds and seed yield/plant. The SCA effects calculated according to Griffing's method equaled the average of SCA effects of each cross and its reciprocal. Meanwhile, in the proposed model, the average of the difference between SCA effects of each cross and its reciprocal equaled the reciprocal effects. This would prove that reciprocal effect provides precise estimation to the interaction effect between nuclear and cytoplasmic genes of the cross and its reciprocal hybrid.

© 2016 Production and hosting by Elsevier B.V. on behalf of Faculty of Agriculture, Ain Shams University. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/

licenses/by-nc-nd/4.0/).

Introduction

Faba bean (Vicia faba L.) is a valuable food legume crop in Egypt and many other Mediterranean countries. Furthermore, this crop can play a key role in sustainable production and management of agriculture and in enhancement total soil

* Corresponding author. Peer review under responsibility of Faculty of Agriculture, Ain-Shams University.

nitrogen fertility of nutrient poor soil through biological atmospheric nitrogen fixation (Lindemann and Glover, 2003).

On the other hand, faba bean is a self-pollinating plant with significant levels of outcross and inter-cross, ranging from 20% to 80% (Suso and Moreno, 1999) depending on tested genotype and surrounding environmental effects. The genetic improvement of crop desired traits depends on the nature and magnitude of genetic variability and interactions involved in the inheritance of these traits. It can be estimated using diallel cross technique, which provides early information on the

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0570-1783 © 2016 Production and hosting by Elsevier B.V. on behalf of Faculty of Agriculture, Ain Shams University. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

genetic behavior of these traits in the first (Fi) generation (Chowdhry et al., 1992). This technique may also result in the production of new genetic combinations performance (negatively or positively), maybe exceeding over the parents. However, the parental superiority may not depend so much on their actual performance as on their ability to combine well and through transgressive segregates (Zhang and Kang, 1997).

The combining ability considers as an important criteria for plant breeders, where it is useful in connection with testing procedures to study and compare the performance of lines in hybrid combinations and the nature of gene action. So, the plant breeders are interesting with the gene effect estimates to apply the most effective breeding procedure for the improvement of the desired attributes. Moreover, the choice of the most efficient breeding methodology mainly depends upon the type of gene action controlling the genetic behavior of most agronomic and economic characters. Nevertheless, for obtaining a clear picture of genetic mechanism of faba bean populations, the total value of variances must be portioned into its genetic components. Hence, exploitation of the genetic components could encourage improving yield potential and its components in faba bean plants, whereas, the superiority of crosses/hybrids over parents for seed yield is associated with manifestation of gene effects in important yield components. These effects may differ from significantly positive to significantly negative for different traits depending on genetic makeup of faba bean parents. The importance of gene action and heritability was previously discussed by Awaad et al. (2005), Darwish et al. (2005), Attia and Salem (2006), El-Hady et al. (2007, 2009), Bayoumi and El-Bramawy (2010), El-Bramawy and Osman (2010, 2012), and Ghareeb and Helal (2014).

Griffing (1956) defined diallel crosses, which have been used extensively in plant breeding. However, general and specific combining ability effects are commonly based on the average effect of the parent when it is used as a female or a male in its hybrid combinations assuming that they are likely to be similar as proposed by Yates (1947). When crosses and their reciprocals are included, the fixed models, only one GCA effect value for each parent and one SCA effect value for each cross combination is estimated. Accordingly, these estimated effects were not separated, showing the contribution of each parent to the cross combination when this particular parent is used as a male or, alternatively, female. The difference between the interaction effect of the cross and its reciprocal is due mainly

to the interaction between the nuclear and the cytoplasmic genes as indicated above. Cytoplasm of the female parent may represent different environment that differs from one parent to another (Ghareeb et al., 2014) and therefore, interacts with nuclear genes differently. Interaction between the nuclear and the cytoplasmic genes was reported by Singh and Brown (1991), Ekiz and Konzak (1991), Maan (1992), and Voluevich and Buloichik (1992).

Partitioning of the general and specific combining ability effects would provide additional information about each parent when it is used as a female or a male in its hybrid combinations (Mahgoub, 2004). Improving the precision of the statistical model used for estimating GCA and SCA effects may provide an effective tool for selecting the breeding method as well as the paired populations to be used in a reciprocal recurrent selection program. In Egypt, on faba bean, no references have been found about the abovementioned research topic.

Therefore, the objectives of the present study were to: (1) compare the GCA and SCA effects before and after partitioning, (2) evaluate the relative contribution of each parent to its cross combination when it is used as a male or a female parent, (3) detect the significant of maternal effects and (4) estimate the relationship between SCA effect and reciprocal effect.

Materials and methods

Genetic materials and cross model

The current investigation was carried out at Giza Research Station, ARC, Giza, Egypt, during two successive growing seasons, 2013/14 and 2014/15. Five faba bean varieties were chosen on the basis of the presence of wide differences among them as shown in Table 1. In 2013/14 season, full diallel crosses (all cross combinations including reciprocals) were made between the five parents. The parents and their 20 F{s (25 genotypes) were grown in 2014/15 season at the 26th November. Each genotype was planted in 4 rows 3 m long, 30 cm apart from one seed spaced at 20 cm. Randomized Complete Block Design with three replications was used under free insect cages. The soil texture of the experimental site was clay loam with pH value of 7.4 and EC of 2.46 dS/m. Cultural practices were applied as recommended for faba bean production in the area to raise a good crop. At harvest, data were recorded on ten individual guarded plants aiming the following

Table 1 Pedigree and special traits of five faba bean parental genotypes.

Genotype Source Pedigree Seed type Characteristics

Giza 3 (Pi) Food Legumes Research Cross (Giza 1 x Dutch Intr. Equina Resistant to foliar disease, high yield

Department 29)

Giza 461 (P2) * FCRI, ARC, Egypt Cross (Giza 3 x ILB938) Equina Resistant to foliar disease, high yield

Nubaria 1 (P3) Single plant selection from Major Recommended for planting in newly

the Spanish cultivar Reina reclaimed lands and resistant to foliar

Blanca diseases

Triple white (P4) Sudan Equina High autofertility, white flower with light

seed coat color and colorless hilum, and

susceptible to insects' storage

Giza 716 (P5) 461/843/83 x 503/453/84 Equina Resistant to foliar diseases and early

maturing

* FCRI (Field Crop Research Institute), ARC (Agriculture Research Center), Egypt.

traits: Plant height (cm), number of branches per plant, number of pods per plant, number of seeds per plant, 100-seed weight (g) and seed yield per plant (g).

Statistical analysis

Analysis of variance was carried out to determine the significance of genotypic differences. When the significant differences among the genotypes were established, the total variance was portioned to genetic factors using two partitioning methods as follows: (1) The diallel analysis model proposed by Hayman (1954) to separate total sum of square into various components, namely, a (additive), b (non-additive, which is further subdivided into b\, b2 and b3), c (maternal) and d (reciprocal differences other than c) and (2) the model suggested by Griffing (1956) method 1 model I was also applied in a modification type, where GCA and SCA effects were partitioned to study the contribution of each parent when it is used as a male or a female in its hybrid combinations, but not on the average performance of male and female parents (Mahgoub, 2011).

Proposed model formula

Griffing's method 1 model I which included parents (P), F^s and their reciprocals was applied. Various effects are estimated according to Griffing (1956) as follows:

Si = 1 2pj (x>- + x-i) -Sij ~ (2) (xij + Xji) ' rij ~ (2) (xij — Xji) •

2p I (Xi• + X• i + Xj• + X j)

Maternal effect is estimated according to Cockerham

(1963) using Griffing's notations as follows: fh — (^p

where x,.: is the sum of the ith female over all males; x ¡: is the sum of the ith male over all females; x/.: is the sum of the jth female over all males; x j is the sum of the jth male over all females; xj: is the mean for the F1 resulting from crossing the ith female and the jth male parents, xji: is the mean for the F1 resulting from crossing the jth female and the ith male parents; g: is the general combining ability effect of the ith parent, bsij: is the specific combining ability effect for the cross between the ith female and the jth male parents (bj — b), is the reciprocal effect involving the ith and jth parents, mi: is the maternal effect of the ith parent and x.. : is the grand total.

For proposed model where GCA effect (b) is partitioned to estimate GCA effect for the parent when it is used as a female in its hybrid combination (bfi), and GCA effect for the same parent when it is used as a male in its hybrid combination (gmi) as follows:

gfi — feW)—fek,

■= (i (X: '

where bfi: is the deviation of the mean performance of the ith parent when it is used as a female, averaged over a set of P males, from the grand mean and bmi: is the deviation of the mean performance of the ith parent when it is used as a male, averaged over a set of P females, from the grand mean where:

9 — Q) (gfi + bmi) and, m — Qj (9 - bmi) -

This proves that the average of the difference between bfi and <bmi is exactly equal to maternal effect (m). In other words, estimation of (bfi — bmi) would provide precise estimation for the maternal effect. General combining ability effect provides estimation for the additive effect. Therefore, maternal effect is mainly additive and expresses how much additive effect is involved.

"Check of computations (gca effect): ^^ b — 0, bfi — 0,

^Cbm — 0 and J2 m — 0.

Specific combining ability effect is partitioned to estimate SCA effect for the cross bsij and for its reciprocal bsji as follows:

2p j ^i• ^ X • i ^ Xj• ^ X :j)

sji - x ji \1p) (xi- ^ x-i ^ xj- ^ xj)

where the average of the partitioned components (bj and j is equal to calculated bj according to Griffing's method, bf is the SCA effect of the ith female and the jth male parent, and bsji is the SCA effect of the reciprocal and the jth female and the ith male parent.

*Check of computations (sca effect): Griffing's £ %

Eby + E b = 2

Reciprocal effect (r) = (|) (bj — bp) and reciprocal effect

rj = —rji.

This proves that the average of the difference between SCA effect of the cross and its reciprocal is exactly equal to the estimated reciprocal effect. Accordingly, this difference provides precise estimation for the reciprocal effect. Testing the significance differences was estimated according to Griffing's method.

Results and discussion

The progress in the breeding program of a certain crop characters depends on the variability in populations and the extent to which the desirable characters are heritable in this respect. However, the knowledge of the genetic architecture of yield and other characters help to formulate a meaningful breeding strategy for developing improved genotypes. Before conducting a complete diallel analysis for all the studied traits, a formal analysis of variance procedure following Steel and Torrie (1980) was carried out to see significant genotypic differences among the studied genotypes because only significant genotypic differences allow further analysis of the data. Therefore, the obtained results and their discussion will be presented in the following.

Diallel analysis

The statistical analysis for faba bean parental genotypes and their crosses revealed presence of highly significant differences among them for all the studied traits, except number of branches/plant (Table 2). These findings provided evidence for the presence of high considerable amount of genetic variability and additive effect (item a) among the parental genotypes and their respective F1 hybrids for all the studied traits

except number of branches/plant. Consequently, complete diallel analysis for all the studied traits except number of branches/plant was done. These results were in harmony with those reported by El-Hosary et al. (1998), Awaad et al. (2005), Attia and Salem (2006), Bayoumi and El-Bramawy (2010), El-Bramawy and Osman (2012), and Ghareeb and Helal (2014). Complete diallel analysis for all the studied traits except number of branches was done.

Genotypic variance was partitioned into various components: additive (a), non additive (b), maternal effects (c) and (d) items (Mather and Jinks, 1971; Aksel and Johnson, 1963). Diallel analysis showed that both additive (a) and non additive (b) components of variance were significant and both are important in genetic control of all traits in F{s. However, the additive component accounted for greater proportion than the non additive component. Significant (b1) values were obtained for all the studied traits except number of pods/plant, revealing that the dominance deviation of the genes is predominantly in one direction. But (b2) was highly significant for all the studied traits, pointing to presence of asymmetrical gene distribution of dominant and recessive alleles, and thus some parents considerably have more number of dominant alleles than others. However, (b3) was highly significant for 100-seed weight, indicating significance of the part of dominance deviation which was not attributable to (bi) and (b2). Number of seeds, 100-seed weight and seed yield/plant recorded significant item (c) meaning the presence of maternal effects, meanwhile plant height, number of seeds and seed yield/plant revealed significant values for item (d), pointing to the presence of reciprocal differences (Singh and Chaudhary 1985).

Significant reciprocal effects in the expression of yield and other important traits have been reported by Chowdhary et al. (2007) in bread wheat and, Topal et al. (2004) in durum wheat. This indicates maternal influence or role of maternal parent in determining the phenotype of F1 and thus importance of selecting the parents while making crosses, and also there exists evidence for expression of heterosis in yield and almost of agronomic traits. Radawan et al. (2010) pointed out the importance of cytoplasmic effects on faba bean; the present effects in the reciprocal crosses indicate extra nuclear factors influencing some traits. This suggests that the recipro-

cal effects may be widely spread in faba bean and that trait expression in F1 hybrid may be due to the function of both genetic and cytoplasmic factors.

Performance of parents and their hybrids

There is no doubt genetically, that the offspring which is produced from different hybrids may display a higher yielding potential compared to the mean yield of its parents. Mean values of the five faba bean varieties and their respective hybrids for the significant traits are shown in Table 3.

The behavior of plant height character was significantly differed from one genotype to another over all faba bean genotypes. The parent values ranged from 135.00 cm for the variety Giza 3 (P1) to 100.00 cm for the variety Triple white (P4). Meanwhile, the values of plant height in the different hybrids ranged from 141.67 cm for the cross (Nubaria 1 x Giza 461) to 91.11 cm for the cross (Giza 3 x Giza 716). Therefore, it can note that the reciprocal cross (Nubaria 1 x Giza 461) had the tallest plants.

The parent Triple white (P4) possessed the lowest values for 100-seed weight (51.13 g) and seed yield per plant (32.33 g). Moreover, the parent Nubaria 1 (P3) gave the highest values of 111.70 g and 64.78 g, respectively for the same traits. The hybrid Nubaria 1 x Giza 461 recorded the highest value for seed yield per plant (119.33 g), while, the cross Triple white x Nubaria 1, Giza 3 x Nubaria 1 and Giza 461 x Nubaria 1 recorded the highest values for 100-seed weight (115.06, 114.68 and 113.83 g, respectively).

The parent Giza 3 (P1) gave the highest values for number of pods per plant (33.00) and number of seed per plant (91.00). Moreover, Giza 716 (P5) showed the lowest values of 16.50 pods and 47.92 seeds per plant. On the other side, the hybrid, (Giza 3 x Giza 461) recorded the highest values for number of pods per plant (51.00 pods) and number of seeds per plant (133.33 seeds).

From the above results of mean performance of the parents and their hybrids, it could be concluded that the hybrids had promising characters which can possess the genetic factors for high yield potential. These results could confirm the possibility of selection for these studied characters through the

Table 2 Significance of mean squares due to different sources of variations for all the studied characters according to Hayman's

model.

S.O.V d.f Plant height No. of No. of No. of 100- seed weight Seed yield/ Plant

(cm) branches/plant pods/plant seeds/plant (g) (g)

Genotypes 24 487.31** 2.64ns 310.79** 1376.51** 920.83** 1351.34**

a 4 1314.87** - 1080.74** 4222.79** 3939.93** 2875.58**

b 10 383.06** - 169.00* 798.64** 367.21** 945.81**

b1 1 938.35** - 201.57 1238.18* 976.55** 3774.15**

b2 4 521.15** - 273.25** 1300.62** 124.38** 1260.63**

b3 5 161.53 - 79.09 309.14 439.61** 128.30

c 4 151.37 - 197.59 818.01* 573.73** 1582.86**

d 6 333.30** - 109.28 814.44* 62.18 856.71**

Pooled 48 82.109 1.95 67.91 262.38 32.82 295.50

a: additive variance component, b: non-additive, which is further subdivided into (b1, b2 and b3), c maternal or cytoplasmic variance and d:

reciprocal differences other than c variance component.

* ** and ns indicate significant, highly significant and insignificant at the 0.05 and 0.01 levels of probability, respectively.

Table 3 The mean performance of parents, F1's and their reciprocals for the significant traits.

Genotypes Plant height (cm) No. of pods/plant No. of seeds/plant 100- seed weight (g) Seed yield/plant (g)

Giza 3 (Pi) 135.CC 33.00 91.00 69.31 63.17

Giza 461 (P2) 116.67 25.00 63.00 70.38 44.30

Nubaria 1 (P3) 1C2.64 i6.83 58.95 111.70 64.78

Triple white (P4) 1CC.CC 29.00 81.00 51.13 32.33

Giza 716 (P5) 1C5.CC i6.50 47.92 85.04 41.47

Parents' mean 111.8б 24.07 68.37 77.51 49.21

Pi X P2 131.67 5i.00 133.33 67.79 90.00

Pi X P3 131.67 28.33 78.67 114.68 90.77

Pi X P4 115.CC 46.67 103.00 72.89 75.10

Pi x P5 91.11 ii.ii 36.89 80.49 29.89

P2 X P3 126.67 27.33 81.00 113.83 57.97

P2 X P4 13C.CC 37.00 103.00 63.52 65.53

P2 x P5 124.17 22.92 71.83 91.20 67.33

P3 X P4 116.67 39.67 98.33 89.16 93.07

P3 x P5 117.78 i6.00 62.44 89.21 56.89

P4 x P5 113.61 17.I1 53.53 88.33 47.33

Cross' mean 119.84 29.71 82.20 87.11 67.39

P2 x Pi 14C.CC 29.67 75.33 72.75 54.40

P3 x Pi 123.33 25.00 76.67 93.00 72.20

P4 x Pi 123.33 31.67 81.00 65.10 53.83

P5 x Pi 12C.83 19.97 65.08 75.18 49.00

P3 x P2 141.67 37.67 98.00 100.33 119.33

P4 x P2 128.33 36.00 96.33 72.52 90.73

P5 x P2 1C6.25 19.92 67.50 91.46 61.17

P4 x P3 111.67 31.00 77.67 115.06 67.23

P5 x P3 11C.CC 17.33 60.50 98.83 59.92

P5 x P4 11C.33 17.94 50.50 75.36 37.19

Reciprocals' mean 121.57 26.62 74.86 85.96 66.50

Grand mean 118.94 27.35 76.50 84.73 63.40

LSD at 0.05 8.б0 7.82 15.38 5.44 16.32

hybrids and their respective parents. Thus it allowed the gate open in the front of plant breeders to build future breeding program for high potential yield in faba bean crop. These findings were in agreement with those reported by El-Hosary et al. (1998), Awaad et al. (2005), Darwish et al. (2005), El-Hady et al. (2007), EL-Harty et al. (2009), Bayoumi and El-Bramawy (2010), Ibrahim (2010), El-Bramawy and Osman (2012), and Ghareeb and Helal (2014).

Combining ability

In diallel crosses system, the obtained information about general and specific combining ability for parents and their hybrids may help breeders to identify the best combiners which may be hybridized to build up favorable fixable genes. The estimates of GCA effects "g" are listed in Table 4, where it differed from one parent to another and from character to other. Partitioning of the GCA effects to estimate male "bmi" and female "bfi" effects revealed that their average values equal calculated value according to Griffing's method "gg". The difference between gf and gm revealed that the GCA effect estimates "g" might underestimate the value of the parent, then values of gfi and gm showed better performance in its hybrid combinations.

The parental genotype Giza 461 (P2) had significant or highly significant positive GCA effects ''g" for all studied characters except 100-seed weight. Therefore, this parent could be good combiner for improving these studied characters. Also, the parent Triple white (P4) showed positive and highly significant values for number of pods per plant (4.16) and number of seeds per plant (6.04). However, the parental genotype Nubaria 1 (P3) was good combiner for 100-seed weight (19.02) and seed yield per plant (11.30). Therefore, the parent Nubaria 1 (P3) could be good source for improving 100-seed weight and seed yield per plant in faba bean crop. Consequently, it could be concluded that previously mentioned parental genotypes can be used in faba bean breeding programs. Similar findings were earlier reported by El-Hosary et al. (1998), Darwish et al. (2005), El-Hady et al. (2007), Ibrahim (2010), and El-Bramawy and Osman (2012).

In the proposed method, partitioning of the GCA effects to estimate male and female effects (gfi and gmi) showed better performance when the parent was used as a female or a male in its hybrid combinations. Data in Table 4 show that the average of bfl and gmi effects for 100-seed weight calculated according to Griffing's method (19.02) overestimated the breeding value of the parent 3 (Nubaria 1) compared with its breeding value when it was used as a female parent (11.95), while it

Table 4 The GCA effects (gi), and the adjusted partitioning of the GCA effects to estimate female (bfi) and male (gmO effects of the tested five parents.

Genotype GCA Plant height (cm) No. of pods/plant No. of seeds/plant 100- seed weight (g) Seed yield/plant (g)

Giza 3 (Pi) gi 5.76** 3.60* 6.70** -6.68** 0.75

gfi 1.95 6.68** 12.08** -3.70** 6.39**

gmi 9.56** 0.52 1.32 -9.66** -4.88*

Giza 461 (P2) gl 7.27** * 3.80* 8.73** -3.31** 6.11**

gfi 8.56** 1.04 2.33 -2.39 -5.49*

gmi 5.98** 6.57** 15.13** -4.23** 17.71**

Nubaria 1 (P3) gi -0.46 -1.75 -1.38 19.02** 11.30**

gfi 1.48 -0.31 2.38 11.95** 17.86**

gmi -2.41 -3.18* -5.14* 26.09** 4.73*

Triple white (P4) gi * -4.04 4.16** 6.04** -10.31** -3.93

gfi -3.55* 1.61 1.41 -6.30** -5.10*

gmi -4.54** 6.71** 10.67** -14.32** -2.75

Giza 716 (P5) gi -8.53** -9.82** -20.09** 1.28 -14.23**

gfi -8.45** -9.01** -18.20** 0.44 -13.65**

gmi -8.60** -10.62** -21.98** 2.12 -14.81**

SE GCA, gi 2.563 2.33 4.58 1.62 4.86

* ** and ns indicate significant, highly significant and insignificant at the 0.05 and 0.01 levels of probability.

underestimated when it was used as a male parent (26.09), revealing much higher gbmi than gbfi effect. Likewise in seed yield per plant trait had higher GCA effects when it was used as a female bfi, rather than a male gm (bfi, 17.86 higher than bmi, 4.73), with an average of 11.30 for both effects according to Griffing's method overestimated the breeding value of the parent 3 (Nubaria 1). This indicated that more favorable alleles were provided by the plants of parent 3 when it was used as a female in crosses. Therefore, the parent 3 (Nubaria 1), where the progeny test is based mainly on the performance of the offspring of the female plants, may be more effective in detecting the high gbfi effects and consequently more effective in improving seed yield per plant. The significance of gf of parent 3 (Nubaria 1) may indicate that some gain from selection is expected if the progeny test was based on the performance of the offspring of the female plants as family selection (Genter and Eberhart, 1974).

Adjusted maternal effects

Table 5 shows the adjusted maternal (m) effects of the tested five parents. Data showed that the average of the difference between gbfi and gbmi is exactly equal to maternal effect, which is based on the average of the females over all associated males. Results revealed significant values for number of seeds, 100-seed weight and seed yield/plant. Giza 461 (P2) recorded the highest significant negative maternal effect value for seed yield per plant (—11.60**). Moreover, Nubaria 1 (P3) and Giza 461 (P2) had significant negative maternal effect value for 100-seed weight (—7.07**) and number of seeds per plant (—6.40*), respectively. Meanwhile, Nubaria 1 (P3) and Giza 3 (P1) had significant positive maternal effect value for seed yield per plant (6.56* and 5.63*, respectively). Also Triple white (P4) recorded (4.01*) for 100-seed weight. The average of the difference between gbfi and gbmi effects is exactly equal to the maternal effect calculated according to Cockerham (1963), which is based on the average of the females over all associated males.

Therefore, partitioning of the GCA effects provided additional information to plant breeders about estimating maternal effect. Estimation of maternal effects, which is based on the average of the females over all associated males, would underestimate maternal effect of some specific cross combinations, which may be more important. Therefore, partitioning of the maternal effects leads to the estimation of the reciprocal effects, and this provides estimation of the maternal effects on a hybrid combination basis rather than on the average of all associated male parents (Mahgoub, 2011; Fan et al., 2014).

Specific combining ability

The SCA effects calculated according to Griffing's method and the partitioned SCA effects are presented in Table 6. Plant height and seed yield per plant revealed that, the reciprocal cross (P3 x P2) had significant and much higher values for SCA effects (15.92** and 38.53**, respectively) after partitioning, compared with its cross (P2 x P3) values (0.92 and —22.84**, respectively). But SCA effects calculated according to Griffing's method assumed that SCA effects are the same (8.42** and 7.85*, respectively) for each cross and its reciprocal (equal the average of cross and its reciprocal SCA effects) and do not show this additional information. Likewise, cross (P1 x P2) had SCA effects with highly significant and much higher values after partitioning (16.25** and 41.40**), compared with their reciprocals (P2 x P1) for number of pods and seeds per plant (—5.08* and —16.60**), respectively. Also, cross (P4 x P3) had significant and much higher SCA effect values after partitioning (21.62**), compared with their reciprocal (P3 x P4) for 100-seed weight (—4.28).

Adjusted reciprocal effects

The SCA effects were different (after partitioning) when a genotype was used as female from those when the same genotype was used as male (Mahgoub, 2011; Fan et al.,

Table 5 The adjusted maternal (m) effects of the tested five parent.

Genotypes Plant height (cm) No. of pods/plant No. of seeds/plant 100-seed weight (g) Seed yield/plant (g)

Giza 3 (Pi) -3.81 3.08 5.38 2.98 5.63*

Giza 461 (P2) 1.29 -2.77 -6.40* 0.92 -11.60**

Nubaria 1 (P3) 1.94 1.43 3.76 -7.07** 6.56*

Triple white (P4) 0.49 -2.55 -4.63 4.01* -1.18

Giza 716 (P5) 0.07 0.80 1.89 -0.84 0.58

SE (GCAi-GCA,) 4.052 3.68 7.14 2.56 7.68

* ** and ns indicate significant, highly significant and insignificant at the 0.05 and 0.01 levels of probability.

Table 6 The SCA effects according to Griffing' s method (upper) and SCA effects according to the proposed method (bold, lower) of

Fi crosses (F1) and their reciprocals (F1r).

Genotypes SCA Plant height (cm) No. of pods/plant No. of seeds/plant 100- seed weight (g) Seed yield/plant (g)

P1, P2 Griffing's 3.87 5.59* 12.40** -4.47 1.94

P1 X P2 F1 -0.30 16.25** 41.40** -6.95* 19.74**

P2 X P1 Fh 8.03** -5.08* -16.60** -1.99 -15.86**

P1, P3 Griffing's 3.27 -2.53 -4.15 6.77 6.04

P1 X P3 F1 7.44** -0.86 -3.15 17.61** 15.32**

P3 X P1 F1r -0.90 -4.20 -5.15 -4.07 -3.25

P1, P4 Griffing's -1.49 4.06 2.77 1.25 4.24

P1 X P4 F1 -5.65* 11.56** 13.77** 5.15 14.88**

P4 X P1 F1Rec 2.68 -3.44 -8.23* -2.64 -6.39

P1, P5 Griffing's -10.20** -5.58* -12.12** -1.50 -10.48**

P1 X P5 F1 -25.06** -10.01** -26.22** 1.16 -20.03**

P5 X P1 Fir 4.67 -1.16 1.97 -4.16 -0.92

P2, P3 Griffing's 8.42** 3.10 5.65 6.64 7.85*

P2 X P3 F1 0.92 -2.07 -2.85 13.39** -22.84**

P3 X P2 F1r 15.92** 8.26** 14.15** -0.11 38.53**

P2, P4 Griffing's 7.00** 1.19 8.40* -3.09 12.55**

P2 X P4 F1 7.83** 1.69 11.73** -7.59* -0.05

P4 X P2 F1r 6.17* 0.69 5.06 1.41 25.15**

P2, P5 Griffing's -2.47 0.08 4.52 8.63* 8.97**

P2 X P5 F1 6.49** 1.58 6.69* 8.50* 12.06**

P5 X P2 F1r -11.43** -1.42 2.35 8.76* 5.89

P3, P4 Griffing's -0.26 5.57* 6.85* 8.67* 9.39**

P3 X P4 F1 2.24 9.91** 17.18** -4.28 22.30**

P4 X P3 F1r -2.76 1.24 -3.49 21.62** -3.53

P3, P5 Griffing's 3.94 0.88 6.44* -11.02 -2.06

P3 X P5 F1 7.83** 0.21 7.41* -15.83 -3.57

P5 X P3 F1r 0.05 1.55 5.47 -6.02 -0.55

P4, P5 Griffing's 5.60* -4.16 -10.43** 6.14 -2.98

P4 X P5 F1 7.24** -4.58* -8.92** 12.63** 2.09

P5 X P4 F1r 3.97 -3.75 -11.95** -0.35 -8.05*

SE SCAy 5.284 4.80 9.44 3.34 10.02

*, ** and ns indicate significant, highly significant and insignificant at the 0.05 and 0.01 levels of probability.

2014). Reciprocal effects (b) calculated according to Griff-ing's method and the partitioned SCA effects are presented in Table 7. Crosses between the parents Giza 461 (P2) and Nubaria 1 (P3) recorded the highest and significant reciprocal effect (b) values for yield per plant (±30.68**). Meanwhile, number of seeds and seed yield per plant showed

the greatest and significant reciprocal effects (±29.00** and ±17.80**) by the progeny of crosses Giza 3 (P1) and Giza 461 (P2), respectively. The highest and significant reciprocal effects (±14.86** and ±14.10**) were obtained in crosses between Giza 3 (Pi) and Giza 716 (P5) for plant height and number of seeds/plant. Then, reciprocal effect values

Table 7 Reciprocal (b) effects calculated according to Griffing's method and adjusted method (as the same values, but in two

directions).

Genotypes Plant height (cm) No. of pods/plant No. of seeds/plant 100-seed weight (g) Seed yield/plant (g)

Pi, Pi -4.17" 10.67** 29.00** -2.48 17.80**

Pi, P3 4.17** 1.67 1.00 10.84** 9.28*

Pi, P4 -4.17** 7.50** 11.00** 3.90 10.63*

Pi, P5 -14.86** —4.43 — 14.10** 2.66 —9.56*

Pi, P3 -7.50** —5.17 —8.50* 6.75** —30.68**

Pi, P4 0.83 0.50 3.33 -4.50* —12.60**

Pi, P5 8.96** 1.50 2.17 -0.13 3.08

P3, P4 2.50* 4.33 10.33* -12.95** 12.92**

P3, P5 3.89** —0.67 0.97 -4.81* —1.51

P4, P5 1.64 —0.41 1.51 6.49** 5.07

SE R^- 6.407 5.82 11.45 4.05 12.15

There are r y values, whereas Tyj had the same values with different sign (ry = — ry).

between (by) Giza 3 (Pj) x Giza 461 (P2) and between (by) Giza 461 (P2) x Giza 3 (P1) had the same values with different sign (by — —bji) (Fan et al., 2014).

In diallel cross model, the occurrence of reciprocal differences for all studied components, indicated that the cytoplasm of maternally inherited factors interacts with nuclear genes to control the response of faba bean genotypes. The reciprocal effects strongly influenced estimates of SCA effects.

Conclusion

The statistical analysis of the five parents and their possible hybrids (F1), revealed highly significant differences for all the studied traits except number of branches per plant. Preliminary information about the presence of significant variation is obtained from Hayman diallel analysis to divide total sum of square into various items, i.e. additive, non-additive components, maternal or cytoplasmic and other reciprocal differences. The cytoplasmic components were significant for number of seeds/plant, weight of 100 seeds and seed yield/plant. General and specific combining ability effects were partitioned according to a proposed model to estimate them for each parent when it is used as a female or a male in its hybrid combinations. Results revealed that estimated GCA effects according to Griffing's method are equal to the average of GCA effects of each parent, after partitioning, when it is used as a male and a female in its hybrid combinations. In addition, the average of the difference between female and male GCA effects would provide valid and precise estimation of the maternal effect as previously confirmed by Hayman analysis for number of seeds/plant, weight of 100 seeds and seed yield/plant. Giza 461 and Nubaria 1 had significant negative maternal effect values for number of seeds per plant and 100-seed weight, respectively. This would prove that maternal effect provides precise estimation to the favorable alleles, which are mainly additive ones. The SCA effects calculated according to Griffing's method are the average of SCA effects of each cross and its reciprocal. Meanwhile, the average of the difference between SCA effects of each cross and its reciprocal, according to the proposed model, is equal to the reciprocal effects. Crosses between the parents Giza 461 and Nubaria 1

recorded the highest and significant reciprocal effect (b) values

for yield per plant.

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