Genetic study of agronomic traits in barley based diallel cross analysis.
Barely is an autogamous plant and one of the oldest members of family Poaseae. This plant is farmed in most provinces of Iran, and has the second rank among other cereals of the world. Because of three reasons namely broad ecological compatibility, its use in human's and animal's food, barely has attracted big attentions . The study of genetic conditions of different agricultural plants is one of the essential factors for the success of inbreeding plans. Therefore it is required that precise and comprehensive information of the genetic parameters controlling the components of yield is collected and used for making decisions on the selection of an appropriate breeding method. Such information is collected using different methods. One method of collecting information is diallel cross method. This method was introduced by Jinks, Jinks and Hayman, and Hayman & Griffing in 1950, and it was completed by Ponny et al. and Wright. Diallel cross is used to study the genetic diversity and polygenic systems of quantitative traits. As the most important traits are inherited in a quantitative manner , therefore the results of such crosses are valuable for the improvement of the traits inside and among populations, as well as the production of cultivars [29,25].
Graphic analysis based on Jinks and Hayman  makes it possible to have access to the information such as average dominant degree, the ratio of distribution and dispersion of dominant and recessive alleles in parents, and the direction of dominance . In 1986, Koroleva studied in a diallel project the seed weight per plant, weight of 1000 barley seeds in [F.sub.1] and [F.sub.2], and in both studied traits gene overdominance was observed. Moreover, the genes had positive effects . Chaudhry et al. studied five cultivars of barley using diallel cross method. The variance analysis of generations showed that the traits plant height, number of seeds per spike, weight of 1000 seeds, and seed yield in plant are controlled mainly in form of overdominance. In the control of spike length inheritance, incomplete dominance and additive effects were observed.
Sing et al.  evaluated the inheritance of seed yield and yield components using generation mean analysis. They found out that additive and non-additive effects are important for the traits number of seeds per spike, 100 seeds weight, and seed yield, and for most traits, there are dual epistasis effects. Nooral et al.  reported that plant height and weight of 1000 seeds of weight are under additive gene effects.
The research conducted by Koval  on the awn length trait shows that the control of this trait by a gene is in form of incomplete dominance. Zaluski et al. 1991) studied the quantitative traits in bread wheat using graphic analysis, and showed that genes control the traits plant height and spike length by incomplete dominance, weight of 1000 seeds by complete dominance, number of seeds per spike, number of spikelet per spike, and seed weight per spike by overdominance. The present research has been carried out to study the genetic control manner of the traits of yield and estimate the value of different genetic parameters in these traits.
Materials and Methods
In this research, seven cultivars of barely including Valfajr, Makoui, Kavir, Reyhan, Gorgan 4, Nosrat, and Nimruz were crossed in form of semi-diallel. The parents and progeny [F.sub.1] were cultivated and evaluated in the research farmland of the Islamic Azad University of Karaj in the agricultural year 2009-2010 in form of complete randomized blocks with three replicates. Over the course of the growth and development, crops were weeded manually and quantitative traits including seed yield, biologic yield, days to heading, spike length, awn length, 100-seed weight, stem diameter, seed length, and peduncle length in five plants selected from each trial plots were studied and noted. The information was analyzed based on complete randomized blocks plan.
Considering the significance of the variance of genotypes, Hayman method  was used to calculate the trait controlling genetic parameters including additive effect, dominance effect, average dominance degree, number of trait controlling genes, the ratio of the genes having positive and negative effects in parents, broad-sense and narrow-sense heritability, and graphic analysis. SAS (for simple variance analysis), Dial98 (for genetic analysis), and Excel (for graphs) were used for the analysis of the obtained data.
Results and Discussion
In table 1, the results of variance analysis obtained based on the method proposed by Hayman  have been shown. Component a, which is an estimation of additive variance has been significant at 1% probability level for all studied traits.
Moreover, component "b", which is related o the difference between hybrids and parents and arising out of the non-additive effects of genes, has been significant at 1% of probability level for all traits. Akram  conducted an analysis using Hayman method to study the role of additive and non-additive effects in the control of traits including seed yield, cluster length, and 100-seed weight. Based on the method proposed by Hayman , this component of variance was divided into [b.sub.1], [b.sub.2], [b.sub.3]. Component [b.sub.1] means the comparison of parents with crosses; in other words, this component indicates average heterosis. Component b1 has been significant for all traits excluding seed length, stem diameter, and spike weight, which means that these traits are controlled by directive dominance. Component [b.sub.2] shows the special heterosis of each parent.
The significance of this component determines if the deviation of [F.sub.1] from the average parents changes from one parent to other parent. This happens when the frequency of dominant allele are different. This component was significant for traits such as spike length, spike weight, awn length, 100-seed weight, stem diameter, and insignificant for other traits.
The component [b.sub.3] is the most part of the dominance, and according to Mather and Jinks , this component is equal to the value of narrow-sense combining ability in the method 3 of Griffing . This component has been significant for all studied traits.
Estimation of Genetic Parameters:
The estimation of genetic parameters has been provided in table 2. The variance analysis of genetic components showed that additive variance (D) has been significant for the traits including seed yield, biologic yield, days to heading, spike length, stem diameter, and 100-seed weight. Dominance variance (H1) has been significant for all studied traits excluding stem diameter and seed length. Moreover, another form of non-additive dominance (H1, H2) indicates the additive and non-additive effects controlling these traits genetically (Table 2). The average degree of gene dominance [(H1/D).sup.1/2] has been < 1 for the traits seed yield, biologic yield, days to heading, and spike length. This amount has been > 1 for spike weight, awn length, 100-seed weight, stem diameter, seed length, and peduncle length. Being greater or lesser than 1 show the relative dominance and overdominance of genes. Thus, it was concluded that the traits seed yield, biologic yield, days to heading, and spike length were controlled by relative dominance of genes, and other traits were affected by the overdominance of genes (table 2).
It is pertinent to mention that this overdominance can be of false type and due to gamete disequilibrium. The overdominance effects on the traits seed yield [21,2], spike weight , and 100-seed weight , as well as incomplete dominance effects on biologic yield , peduncle length , spike length  have been reported previously. The value of [h.sup.2] has been significant for the traits seed yield, and biologic yield. This shows the high degree of dominance effects of heterozygote gene locations.
The ratio of H2/4H1 indicates the symmetry of the frequency of dominant and recessive alleles in all gene location controlling traits . This ratio is less than 0.25 for the traits including peduncle length, stem length, spike length, spike weight, awn length, and 100-seed weight. This amount of ratio indicates the higher frequency of recessive allele for the said traits. This ratio has been closely around 0.25 for days to heading, which indicates the equal frequencies of dominant and recessive allele for this trait. The value of genetic ratio H2/[h.sup.2] estimated for seed yield, and biologic yield indicates the involvement of at least four genetic groups and for other traits, this ratio shows that there has been at least one genetic group involved in the control of heredity (table 2). The symbol (mf1-mp) indicates the direction of dominance, and for all traits, this symbol has been positive, indicating that reductive alleles have been dominant (table 2).
The value of KD/KR shows the ratio of genes with dominance effects to genes with additive effects. In case this ratio is greater than 1, it indicates that gene locations with dominance effects are more frequent than those with additive effects. In case this value is less than 1, it indicates that gene locations with additive effects are more frequent that those with dominance effects. According to the results shown in table 2, the value of KD/KR, which is the ratio of genes with dominance effects to the genes with additive effects in parents for the traits including days to heading, spike length, awn length, and 100-seed weight, is greater than 1; this shows that the genes with dominance effects are more frequent than those ones with additive effects.
For other traits, this ratio was less than one, which indicates that genes with additive effects have been more frequent than those with dominance effects. The algebraic symbol F confirms this result (Table 2).
The least degree of general heritability equal to 70% has been reported for peduncle length and the most degree of the same equal to 88% has been reported for spike length. The higher value of broad-sense heritability shows that yield components are under control of genetic factors (table 2). The values of narrow-sense heritability vary from 35% for awn length to 72% for spike weight and seed yield. Considering the broad difference of broad-sense heritability and narrow-sense one related to the traits awn length, seed diameter, and peduncle length, it can be claimed that dominance effects are more controlling than additive effects of genes are (table 2). The calculation of heritability is important because it can provide us with the information required for the transfer of traits from parents to their progeny, accelerates the evaluation of genetic and environmental effects on phenotype diversity, and helps selection. Mostafavi et al.  reported the highest value of broad-sense heritability for awn length and spike weight that were equal to 0.95 and 0.76 respectively. Moreover, the highest value of narrow-sense heritability equal to 0.71 was reported for awn length.
The regression coefficients of variance and covariance of lines respectively has and lacked significant different with zero and one. This hypothesis was confirmed for all traits excluding seed length by deleting Nosrat parent, and spike weight by deleting Nimruz parent, deviated both from regression line. The cross of regression line from coordinates' center indicates that there is complete dominance.
In case regression line crosses Wr axis in the upper part (positive section) or lower part (negative section) of coordinate center, it shows relative dominance and gene overdominance are active. According to the abovementioned explanations, regression line Wr over Vr crossed Wr axis in the positive section for the traits seed yield, biologic yield, days to heading, spike length, spike weight, stem diameter, seed length, and peduncle length. This shows that relative dominance is more effective in the genetic control of these traits. For other traits including awn length, and 100-seed weight, the regression line crossed Wr axis in the negative section; this indicates that the overdominance effects of genes are here effective in the control of these traits.
The distribution manner of parents along the regression line shows in turn the ratio of the frequency of dominant and recessive genes. It means that those parents that are closer to the crossing of regression line and Wr axis have more dominant genes, and any one farther have higher percent of recessive genes. That is because the recessive homozygote parent has higher variance and covariance; as a result, it is place over the regression line. It is understood that the crossing of these genotypes may lead to the formation of appropriate hybrids.
According to the abovementioned explanations, the closest and furthest cultivars to the origin of coordinates were as follows: Nosrat and Reyhan for seed yield, biologic yield, and awn length; Gorgan 4 and Reyhan for days to heading and spike weight; Gorgan 4 and Kavir for spike length and 100-seed weight; Gorgan 4 and Valfajr for stem diameter; Gorgan 4 and Valfajr for seeld length; Valfajr and Gorgan 4 for seed length, and Reyhan and Kavir for peduncle length.
Therefore, it can be expected that the crossing of Nosrat and Reyhan cultivars leads to the production of products with high values of the traits seed yield, biologic yield, and hybrid awn length, since the combination of these cultivars lead to higher degree of heterosis. For other traits, we can forecast as mentioned.
In general, according to the results of this research, seed yield and yield components are controlled by additive and non-additive effects of genes. The significant additive and non-additive variance of the studied traits confirmed the role of additive and non-additive effects in the control of these traits. Winder and Lebsack  showed that the production of maximum seed yield is possible in Durum wheat, and this is only possible by exploiting two additive and non-additive effects of genes.
In conclusion, it is recommended that in case the main part of genetic variance of any trait is due to the additive effects of the gene, selection can play a great role in breeding method of that trait. Moreover, if the non-additive effects of the genes played a considerable role in some traits, selection under self-fertilization conditions is not recommended. Thus, it is suggested that hybridization method and selection in the segregating generations to be used which can be more efficient.
[FIGURE 1 OMITTED]
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(1) Seyed mohammad mehdi seyed aghamiri, (2) Khodadad mostafavi, (2) Abdollah mohammadi
(1) Graduated M. Sc. Student of Plant Breeding, Karaj Branch, Islamic Azad University, Karaj, Iran
(2) Department of Agronomy and Plant Breeding, Karaj Branch, Islamic Azad University, Karaj, Iran
Seyed mohammad mehdi seyed aghamiri, Graduated M. Sc. Student of Plant Breeding, Karaj Branch, Islamic Azad University, Karaj, Iran
E-mail: firstname.lastname@example.org, Tel: +989365686610
Table 1: Analysis of variance of a 7x7 half-diallel cross set for ten characters agronomic in barley (according to Hayman 1954). S.O.V df MS YLD BIY DSP block 2 [1326.06.sup.ns] [10473.75.sup.ns] 90.73 ** a 6 114208 ** 887784 ** 246.75 ** b 21 9497.56 ** 75025.96 ** 48.82 ** b1 1 118292.9 ** 936381.8 ** 276.68 ** b2 6 [1135.74.sup.ns] [10103.87.sup.ns] [22.15.sup.ns] b3 14 5310.11 * 41324.31 * 43.98 ** Error 2729.35 22149.61 15.94 S.O.V MS SPL SPW AWL W100 S block 0.99ns 1.84 ** 20.29 ** [0.17.sup.ns] a 22.07 11.94 20.13 ** 3.36 ** b 2.50 ** 0.94 ** 7.42 ** 0.58 ** b1 4.26 ** 0.07 ns 38.04 ** 2.15 ** b2 1.75 ** 1.61 ** 7.26 ** 0.45 ** b3 2.70 ** 0.72 ** 5.31 ** 0.52 ** Error 0.52 0.29 1.98 0.11 S.O.V MS STD SEL PDL block 2.72 ** [0.05.sup.ns] 50.73 ** a 3.75 ** 3.53 ** 100.79 ** b 0.34 ** 0.59 ** 28.85 ** b1 0.17 (tm) [0.19.sup.ns] 136.96 ** b2 0.49 ** [0.13.sup.ns] [17.68.sup.ns] b3 0.29 * 0.81 ** 25.92 ** Error 0.14 0.21 9.64 *, ** Significant at the 0.05, and 0.01 probability levels, respectively. a, additive effect; b, dominance effect; b1, mean dominance deviation; b2, dominance deviation due to each parent; b3, dominance deviation due to each crossing combination. YLD: grain yield, BIY: biological yield, DSP: days to heading, SPL: spike length, SPW: spike weight, W100S: 100 seed weight, STD: stem diameter, SEL: seed length, PDL: peduncle length. Table 2: Genetic parameters for ten agronomic traits in barley MS Parameters YLD BIY Mp 474.01 1337.21 Mf1 555.08 1565.29 D [+ or -] [+ or -] 2275.8 ** [+ or -] 17690.8 ** S.E( D) 7129.4 52603.6 H1 [+ or -] [+ or -] 1808.7 * [+ or -] 15169.2 * S.E( H1) 4132.9 32383.8 H2 [+ or -] [+ or -] 1599.8 ** [+ or -] 13395.4 * S.E (H2) 4549.5 35552.3 F [+ or -] - [+ or -] 1781.5 * - [+ or -] 14085.5 * S.E (F) 3866.9 32705.7 [h.sup.2] [+ or -] 5785.6 ** [+ or -] 48363.5 ** [+ or -] S.E 18899.3 149250.8 ([h.sup.2]) E [+ or -] [+ or -] 127.4 ** [+ or -] 1046.8 ** S.E (E} 909.8 7383.2 [H.sub.2]/ 0.28 0.27 4[H.sub.1] [([H.sub.1/D). 0.761 0.785 sup.1/2] KD/KR 0.47 0.43 K 4.1 4.2 (mf1-mp) 81.07 228.08 [h.sub.b] 0.87 0.87 [h.sub.n] 0.72 0.71 MS Parameters DSP SPL Mp 158.19 8.29 Mf1 162.11 8.77 D [+ or -] [+ or -] 11.8 * [+ or -] 0.52 ** S.E( D) 27.9 2.02 H1 [+ or -] [+ or -] 11.29 * [+ or -] 0.49 ** S.E( H1) 23.4 1.61 H2 [+ or -] [+ or -] 8.89 * [+ or -] 0.36 ** S.E(H2) 22.1 1.33 F [+ or -] [+ or -] [12.79.sup.ns] [+ or -] [0.56.sup.ns] S.E(F ) 7.4 0.27 [h.sup.2] [+ or -] [23.39.sup.ns] [+ or -] [0.5.sup.ns] [+ or -] S.E 42.8 0.62 ([h.sup.2]) E [+ or -] [+ or -] 0.77 ** [+ or -] 0.03 ** S.E(E} 5.3 0.17 [H.sub.2]/ 0.24 0.21 4[H.sub.1] [([H.sub.1/D). 0.916 0.893 sup.1/2] KD/KR 1.34 1.16 K 1.9 0.47 (mf1-mp) 3.92 0.48 [h.sub.b] 0.75 0.88 [h.sub.n] 0.50 0.66 MS Parameters SPW AWL Mp 2.96 11.75 Mf1 3.03 13.20 D [+ or -] [+ or -] [0.15.sup.ns] [+ or -] [1.06.sup.ns] S.E (D) 0.18 1.81 H1 [+ or -] [+ or -] 0.27 ** [+ or -] 1.77 ** S.E (H1) 0.75 4.89 H2 [+ or -] [+ or -] 0.16 ** [+ or -] 1.22 ** S.E (H2) 0.44 3.66 F [+ or -] [+ or -] 0.15 ** [+ or -] [1.57.sup.ns] S.E (F) 0.61 1.34 [h.sup.2] [+ or -] [0.08.sup.ns] [+ or -] [3.01.sup.ns] [+ or -] S.E 0.03 5.91 ([h.sup.2]) E [+ or -] [+ or -] 0.01 ** [+ or -] 0.09 ** S.E (E} 0.09 0.66 [H.sub.2]/ 0.15 0.19 4[H.sub.1] [([H.sub.1/D). 2.038 1.645 sup.1/2] KD/KR 10.76 1.58 K -0.07 1.6 (mf1-mp) 0.07 1.45 [h.sub.b] 0.87 0.72 [h.sub.n] 0.72 0.35 Parameters W100S STD Mp 5.71 3.93 Mf1 6.05 4.03 D [+ or -] 0.24 [+ or -] 0.08 ** S.E (D) 0.17 H1 [+ or -] [+ or -] 0.11 ** [+ or -] [0.11.sup.ns] S.E (H1) 0.39 0.21 H2 [+ or -] [+ or -] 0.08 ** [+ or -] [0.07.sup.ns] S.E (H2) 0.31 0.13 F [+ or -] [+ or -] [0.1.sup.ns] - [+ or -] [0.1.sup.ns] S.E (F) 0.006 0.08 [h.sup.2] [+ or -] [0.17.sup.ns] [+ or -] [0.06.sup.ns] [+ or -] S.E 0.33 0.01 ([h.sup.2]) E [+ or -] [+ or -] 0.01 ** [+ or -] 0.01 ** S.E (E} 0.038 0.05 [H.sub.2]/ 0.20 0.16 4[H.sub.1] [([H.sub.1/D). 1.285 1.109 sup.1/2] KD/KR 1.01 0.65 K 1.1 0.04 (mf1-mp) 0.34 0.1 [h.sub.b] 0.86 0.81 [h.sub.n] 0.57 0.67 MS Parameters SEL PDL Mp 10.02 27.33 Mf1 10.12 30.09 D [+ or -] [+ or -] [0.12.sup.ns] [+ or -] [4.82.sup.ns] S.E (D) 0.23 6.68 H1 [+ or -] [+ or -] [0.14.sup.ns] [+ or -] 7.01 * S.E (H1) 0.23 14.73 H2 [+ or -] [+ or -] 0.11 * [+ or -] 5.28 * S.E (H2) 0.25 12.94 F [+ or -] - [+ or -] [0.12.sup.ns] - [+ or -] [5.94.sup.ns] S.E (F) 0.11 0.08 [h.sup.2] - [+ or -] [0.08.sup.ns] [+ or -] [12.17.sup.ns] [+ or -] S.E 0.001 20.89 ([h.sup.2]) E [+ or -] [+ or -] 0.01 ** [+ or -] 0.47 ** S.E (E} 0.069 3.21 [H.sub.2]/ 0.27 0.22 4[H.sub.1] [([H.sub.1/D). 1.009 1.484 sup.1/2] KD/KR 0.63 0.99 K -0.01 1.6 (mf1-mp) 0.1 2.76 [h.sub.b] 0.75 0.70 [h.sub.n] 0.54 0.39 Mp (Mean of parent), MF1 (Mean of F1s), KD-KR = [(4DH1)1/2 + F] - [(4DH1)1/2-F], K = [h2/H2]. *, ** Significant at the 0.05, and 0.01 probability levels, respectively. [beta]-1 (regression coefficient), a (intercept), D [+ or -] S.E. (D) (additive variance), H1 [+ or -] S.E. (H1) (dominance variance), H2 [+ or -] S.E. (H2) (dominance variance), F [+ or -] S.E. (F) (product of add. by dom. effects), h2 [+ or -] S.E. (h2) (square of difference Pvs All), E [+ or -] S.E. (E) (environmental variance whole), (H1-D)1/2 (average of degree dominance), H2-4H1(balance of positive and negative alleles), KD-KR (proportion of dominance genes), K (number of effective factors), MF1-Mp (direction of dominance), [h.sub.b] (broad sense heritability), [h.sub.n] (narrow sense heritability). YLD: grain yield, BIY: biological yield, DSP:days to heading, SPL: spike length, SPW: spike weight, W100S: 100 seed weight, STD: stem diameter, SEL: seed length, PDL: peduncle length.
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|Title Annotation:||Original Article|
|Author:||aghamiri, Seyed mohammad mehdi seyed; mostafavi, Khodadad; mohammadi, Abdollah|
|Publication:||Advances in Environmental Biology|
|Date:||Jan 1, 2012|
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