# Contextualization of Trait Nexus and Gene Action for Quantitative and Qualitative Characteristics in Indian Mustard

Authors: Sumanta Prasad Chand; Sandip Debnath; Mehdi Rahimi; Mohd. Shaikhul Ashraf; Pankaj Bhatt; Saima Ahmed Rahin
Journal: Journal of Food Quality
 (2022)

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## Abstract

After soybean and palm oil, the world’s third largest oil seed crop is rapeseed mustard. Out of the seven consumable oil seed crops grown in India, rapeseed mustard is responsible for one-third of production. Mustard aphid (Lipaphis erysimi Kalt) is considered the primary pest causing mayhem in crop production. Understanding the genetics behind resistance will aid breeders in developing a resistant/tolerant strain. Appropriate parent selection and analyzing gene action contribute to economic benefit maximization. The aims of the current study are to use seven lines and five testers to determine the best-performing parents and crosses based on their general and specific capacity to combine and to examine the level of heterosis for yield and related features like mustard aphid resistance. Due to the self-pollination nature of Indian mustard, Kempthorne’s line X tester method is helpful to judge the combining ability. Therefore, seven lines and five testers of Indian mustard (Brassica juncea L.) were employed in the present study. The findings suggested that there was substantial genetic variation for all traits examined. The mean oil yield of R1 B2-26 × R1 B2-25, JD6 × R1 B2-25, and JD6 × R1 B2-29 hybrids was more significant than that of the ancestors. The results show that R1 B2-26 × R1 B2-25, JD6 × R1 B2-25, and JD6 × R1 B2-29 hybrids produced more oil than their parents. The variance explained by SCA was greater than that explained by GCA, as indicated by the Ϭ2 gca/Ϭ2 sca ratio being less than one for all characters, implying that nonadditive gene actions such as dominance, epistasis, and other interaction effects played an important role in the presence of these attributes. Punjab Local was discovered to be an excellent general merge for reducing crop duration, whereas JD6 was an excellent combiner for seed yield per crop, aphid infestation indicator, seed yield per crop, and oil production per plant. The predictability ratio was found to be less than 0.5 for almost all traits, denoting that the nonadditive gene measure is involved in controlling the nature except days to 50% flowering, aphid infestation index, oil content, and oil yield per plant. Thus, based on these four traits, selection for superior plants may be practiced in later generations.


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## Body

## 1. Introduction

After cereals, oil seeds are the most important crop in India’s agricultural economy. In India, oil seeds are a significant class of economic crops. Oil obtained from them is an essential component of nutrition and is utilized as a starting good for a variety of products, including paints, coatings, hydrogenated oil, soaps and detergents, perfumes, and lubrication, among others. Additionally, the oil seed sector contributes significantly to international trade by exporting oil seed extracts and oil cakes. Brassicas are India’s second largest growing crop (in terms of coverage), accounting for around 9.12 million tonnes (27.29% of total annual vegetable oil seed production) and occupying approximately 6.78 million ha (25.07% total area for oil seed) of farmland [1]. Plant species belonging to the genus Brassica and family Brassicaceae are known as oil seed brassicas. Some popular Indian mustards involve (Brassica juncea (L.) Czern. & Coss.), toria (B. rapa L. var. toria), yellow sarson (B. rapa L. var. yellow sarson), brown sarson (B. rapa L. var. brown sarson), gobhi sarson (B. napus L.), karan rai (B. car (Eruca sativa Mill.). India is one of the world’s leading producers of rapeseed mustard, ranking first in terms of area and second in terms of output after China. Unfortunately, in comparison with other nations, mustard productivity is quite low. Insects are significant biological restrictions that represent a serious danger to mustard from sprouting to maturity, with over 50 insect species reported infesting rapeseed mustard in India, with about a handful of species regarded as critical pests. L. erysimi, Brevicoryne brassicae L., and Myzus persicae Sulzer are some of the aphids that harm rapeseed mustard in India. L. erysimi is one of the principal restricting factors, producing major output losses and a decline in oil content [2, 3]. It decreases yield by 35.4 to 96%, seed weight by 30%, and oil production by 2.75% [4, 5]. In general, the yield losses due to aphid infestation are very crucial. As a result of aphid infestation, yield losses may range from 5.7 to 28.8% as observed by Kumar et al. [6–8]. Genetic resistance is the most economical and environmentally benign approach to managing insects in crops [9]. Under analogous climatic conditions and insect infestation levels, genetic resistance refers to a genotype’s ability to generate better yields of high-quality food than susceptible cultivars. As a result, resistance is quantified in terms of susceptible cultivars. The scope of this paper is to identify the traits that could help choose the right plant parents to generate resistance against aphids.Dreyer and Campbell [10] emphasized the importance of host plant resistance in biological aphid management and biotype establishment. However, resistance to Brassica aphids is considered polygenic in nature [11]. Given the aforementioned varietal resistance, the fundamental tool of integrated pest management is crucial for managing mustard aphids. A better understanding of resistance will enable breeders to create strains that are resistant for an extended length of time. The notion that heterosis breeding produces superior crop varieties is well known. To evaluate numerous possible breeding procedures, a full evaluation of the likelihood for quantitative traits to combine the inheritance and the phenomenon of hybrid vigor is required [12]. To maximize economic returns through the inclusion of desirable traits, knowledge of combing ability is crucial since it elucidates parental selection and the type of gene functioning involved. Because the foundation (parental genotype and phenotype) must be solid enough to carry the burden of building (successful breeding) placed above, parental selection can make or break economic gains. Given the self-pollination nature of Indian mustard, Kempthorne’s [13] approach for determining combining ability is critical for speedily screening lines [14]. It is important to select parents that can combine fruitfully and produce potential segregation in the following generations [15]. Jenkins and Brunson [16] recommended testing lines with open-pollinated cultivars. Sprague [17] calculated inbreeds’ top-crossing ability. Sprague and Tatum, on the other hand, invented the concept of combing ability [18]. The general combining ability of the line is defined as a divergence from the mean of all crossovers. As a result, any given cross has an estimated value equal to the sum of its two parental lines’ general combining skills. The cross, on the other hand, may differ from the estimated value to a larger or small degree. This deviation is called the specific combining ability of the two lines in combination [19]. Specific combining ability is described as the variation between hybrid combination efficiency and what would be expected based on the parental inbred lines’ general combing ability, while general combing ability is mostly caused by additive effects. Specific combining ability is used to find cross pairings that fare better, whereas general combing ability is often used to assess an inbred line’s contribution to hybrid performance [20]. The variation in forecasted combinations that is dictated by the total average performance of the lines involved is referred to as specific combining ability (SCA). As a result, each hybrid/cultivar development breeder must begin with a review of breeding stock for general and specific seed yield and yield component combinability. It is also critical to discover what kind of gene action influences the inheritance of particular yield-influencing attributes [21]. To overcome yield capping, hybridization is used to recombine genes from genotypically diverse parents. Among the many technological choices available to geneticists and plant breeders, heterosis propagating in crops has been the most victorious. Heterosis has been shown to boost yield and other beneficial features, making it a feasible option for improving Indian mustard production [22]. To successfully exploit heterosis, one must identify hybrids that outperform the parent or standard control cultivar. Brassica rapa var. brown sarson was the first described heterosis among brassicas [23]. The following research has established the amount of heterosis in seed yield. Several studies have indicated substantial heterosis accounting for about 13–91% in B. juncea [24–26]. Moreover, hybrids among genetically distant collections showed more heterosis than hybrids within groups [22]. Within this context, the current research sought to identify the best-performing parents and hybrids based on their general and specific ability to combine, as well as to evaluate the level of heterosis for yield and associated traits, such as resistance to mustard aphids, in seven lines and five testers. The basic aspect of this study is to find characteristics that can aid in selecting the best plant parents for aphid resistance.This paper is organized as follows: Section1 is an introduction to mustard plants, aphids, and plant breeding; Section 2 presents the material and methodology followed for our proposed research; Section 3 describes the experimental settings and data sets collected during the experiment and discusses the experimental results; and finally, Section 4 concludes this paper.


## 2. Materials and Methods

The current experiment was conducted at the Institute of Agriculture, Visva-Bharati University’s Agricultural Farm during theRabi season of 2019–2020. Plant height (cm), number of primary and secondary branches per plant, number of siliquae on the main raceme, and number of siliquae on branches were among the twenty-four phenotypic characteristics studied—the number of siliquae on a plant, the number of seeds on each siliqua, and the length of each siliqua are all factors to consider (cm), days until the first flowering, days until half-flowering, and days until maturity, harvest index, aphid infestation index, oil content (percent), phenol content (mg/g), sugar content (mg/g), protein content (percent), yield per plant (g), oil content (percent), and oil yield per plant (g). The experimental material consists of twelve most common Indian mustard genotypes. Seven genotypes were utilized as females: Kranti, JD6, R1 B2-5, Varuna, R1 B2-26, T 6342, and Punjab Local; and five genotypes were used as males: R1 B2-23, R1 B2-8, R1 B2-6, R1 B2-29, and R1 B2-25, designated as testers. Kranti, JD6, Varuna, and T 6342 were released varieties, whereas Punjab Local, R1 B2-5, R1 B2-26, R1 B2-23, R1 B2-8, R1 B2-6, R1 B2-29, and R1 B2-25 were germplasm obtained at the Nalhati Zonal Adaptive Research Station. T 6342 is a national check variety for mustard aphid resistance. The testers exhibit some resistance to mustard aphids but are low yielding. These twelve parents were crossed to develop 35 F1 hybrids using Kempthorne’s line tester mating scheme [13]. F1 seeds were sown in the field alongside their parents in a randomized complete block pattern with three replications within a 3 m × 2 m plot size with a 30 cm × 10 cm plot spacing following recommended agronomic practices. Ten randomly tagged plants per plot per genotype were taken as samples between the 10th and 15th February every year at the initiation of the flowering stage when aphid infestation was at its peak and evaluated for different biochemical parameters [6]. The phenol and sugar content of the inflorescence was determined. The oil and protein content of seeds harvested from ten different plants was assessed using near-infrared reflectance spectrophotometers (NIRs) [27]. The phenol and sugar contents were determined following Singleton et al. [28] and Sadasivam and Manickam [29] protocols and are presented as mg/g dry weight. Data were recorded for damage scoring to calculate the aphid infestation index (AII) as per Bakhetia and Sandhu [30] and Mukherjee et al. [31]. Analyses of variance were presented on the data collected in accordance with Nadarajan et al. [32] to find important differences between genotypes. Combining capacity effects are extremely essential genetic factors for determining the breeding program’s next phase. The variations for general and specific merge abilities were compared with their corresponding error variations, which were evaluated using the analysis of variations for the various traits (Table 1):Table 1
ANOVA for line× tester analysis.
Source of variationDegree of freedomMean squareReplication (r)(r−1)Genotypes (g)(g−1)MS2Parents (p)(p−1)Parents vs crosses1Crosses (c)(c−1)Lines (l)(l−1)MlTesters (t)(t−1)MtLine× testers(l−1) (t−1)Ml×tError(r−1) (t−1)MS1(1)covariance of half−sib of line=Cov.H.S.line=Ml−Ml×trt,covariance of half−sib of tester=Cov.H.S.tester=Mt−Ml×trl,covariance of full sib=Cov.F.S.=Ml−Me+Mt−Me+Ml×t−Me3r+6rCov.H.S.−rl+tCov.H.S.3r.CovHS (average) is calculated by the following formula:(2)CovH.S.average=1r2lt−l−t×l−1Ml+t−1Mt−Ml×tl+t−2.Assuming no epistasis, variance due to GCA (σgca2) and variance due to SCA (σsca2) are calculated as follows:(3)σgca2=Cov.H.S.=1+F4×σA2,σsca2=1+F22×σD2.In addition, dominant genetic variations (σA2 and σD2) were calculated by taking inbreeding coefficient (F) equal to one; that is, F = 1 because both lines and testers were inbred.Thet-test was used to determine the significance of general combining capacity and specific combining capacity effects. Economic or standard heterosis (SH) is defined as the F1’s increased vigor relative to the commercial standard check. It was calculated using mean values, and the t-test was used to determine its significance [33]. After deriving the variance components, the narrow sense heritability was estimated [34]. The ratios (σgca2/σsca2), σA2/σD2, and σA2/σA2+σD2 were employed to estimate the relatable importance of additive versus nonadditive gene actions [35].


## 3. Results and Discussion

### 3.1. Genetic Variability among Parents and Hybrids

In comparison with the parents, hybrids were of short duration and taller; produced more primary and secondary branches per plant; had a greater number of siliquae on the main raceme, siliquae on branches, and siliquae per plant; and had a greater weight, oil content, and seed yield per plant. Among the parents, JD6 produces the most oil, while R1 B2-5 has the shortest duration (Table2). In relation to testers, the lines were shorter, with more test weight, and contained a greater quantity of siliquae per plant. The hybrids’ mean values for siliqua length, first flowering day, days to 50% flowering, the yield of the top 10 cm of the main raceme, biological yield per crop, stick yield per crop, aphid infestation content, and phenol content were all within the limits of the parental means. T 6342 × R1 B2-23, R1 B2-26 × R1 B2-29, and Varuna × R1 B2-23 cross-combinations all yielded more oil than the parents on average. Seed number per siliqua, siliqua length, days to first flowering, and phenol content were all same in parents and hybrids. Although genetic diversity and parents’ and hybrids’ mean performance were important parameters for genotypic evaluation, parents with high mean values may not pass this trait onto their hybrids. Using general combining ability (GCA) and specific combining ability (SCA), the effects of these parental and hybrid qualities were calculated.Table 2
Mean performance of lines, testers, and hybrids for yield and its attributed traits in mustard.
GenotypesPHPRBSEBSMRSBTOSSDSSLTWDFIFDFFDMMDYTTBIYSYHIAIIOICPHCSUCPRCYPPOYPLinesKranti154.48.48.147.7131.8179.512.85.04.231.037.2108.077.03.448.834.70.293.437.75.79.720.114.15.3JD 6160.65.75.551.1187.4238.512.73.74.128.734.7113.484.83.850.134.90.313.039.65.89.519.916.36.4R1 B2-5160.65.95.646.9106.5153.313.54.23.831.636.6105.273.63.255.542.70.232.039.05.69.820.112.95.0Varuna131.26.76.548.7128.8177.513.04.44.030.734.7108.377.73.452.238.70.263.139.95.79.720.113.55.4R1 B2-26170.86.66.353.7177.8231.414.54.54.328.732.7109.680.93.848.431.80.351.938.55.99.620.116.66.4T 6342172.56.76.449.7191.9241.613.14.54.330.734.7106.375.63.447.933.60.322.036.25.89.920.314.35.2Punjab Local205.77.97.849.1220.2269.311.44.44.130.834.6123.592.73.026.815.60.452.936.95.49.018.011.24.2Mean (Lines)165.16.86.649.5163.5213.013.04.44.130.335.0110.680.33.447.133.10.322.638.35.79.619.814.15.4TestersR1 B2-23172.57.26.940.5162.6203.113.15.54.235.640.9104.669.02.226.418.20.312.039.85.510.420.88.23.2R1 B2-8170.56.56.838.6105.0143.612.84.63.837.041.0114.777.82.026.918.90.302.034.75.19.819.08.12.8R1 B2-6163.34.44.432.7144.9177.613.94.03.943.548.0110.867.41.322.616.40.281.934.94.910.219.86.22.2R1 B2-29213.55.85.735.1214.9250.012.13.03.940.545.0113.873.21.527.521.10.231.937.04.910.019.76.42.4R1 B2-25192.36.46.440.2203.0243.211.24.03.936.340.8114.478.12.226.517.40.341.937.15.110.019.49.13.4Mean (Testers)182.46.16.037.4166.1203.512.64.24.038.643.1111.773.11.826.018.40.291.936.75.110.119.87.62.8HybridsKranti × R1 B2-23184.710.07.650.5352.6403.111.85.24.433.738.7105.371.73.646.129.70.362.136.65.39.419.516.46.0Kranti × R1 B2-8190.09.57.247.8348.7396.512.84.04.135.740.7106.070.33.360.245.40.252.335.55.59.619.614.95.3Kranti × R1 B2-6200.09.510.249.5354.9404.412.74.94.234.739.7105.771.03.457.141.50.272.238.55.49.719.515.56.0Kranti × R1 B2-29188.59.57.350.2361.0411.212.84.44.333.338.7105.372.03.646.129.80.352.138.95.49.419.516.36.3Kranti × R1 B2-25197.79.58.753.2379.9433.012.84.44.432.337.3105.373.02.939.721.70.452.040.25.29.219.418.07.2JD 6 × R1 B2-23190.06.36.737.1386.6423.712.33.94.432.337.3105.373.03.952.734.60.342.037.75.29.219.418.06.8JD 6 × R1 B2-8185.36.43.750.7356.1406.812.14.14.233.738.7105.071.33.550.734.50.322.135.95.39.419.516.25.8JD 6 × R1 B2-6185.09.39.253.5376.7430.212.24.04.231.737.0105.373.73.941.323.30.442.038.75.29.219.418.07.0JD 6 × R1 B2-29174.99.69.653.7379.2432.913.04.14.231.735.7105.774.04.058.940.50.312.040.05.39.219.418.47.4JD 6 × R1 B2-25194.09.610.153.6385.6439.313.34.44.331.335.3105.073.73.045.827.40.402.040.45.39.119.418.37.4R1 B2-5 × R1 B2-23181.27.66.647.5336.5384.012.64.53.736.040.0105.069.03.238.824.30.372.339.55.59.619.614.55.7R1 B2-5 × R1 B2-8176.26.96.446.5326.2372.712.84.14.036.340.3104.368.03.136.322.30.392.337.45.59.619.614.05.2R1 B2-5 × R1 B2-6170.07.56.649.0375.2424.212.14.14.134.738.7105.070.33.436.821.40.422.239.75.49.519.615.46.1R1 B2-5 × R1 B2-29177.58.97.050.5364.4414.911.74.24.234.338.3105.070.73.538.722.40.422.139.85.39.419.516.36.5R1 B2-5 × R1 B2-25181.59.36.653.5378.1431.611.74.24.432.036.0105.073.02.938.820.60.472.040.85.39.219.418.17.4Varuna × R1 B2-23159.97.87.553.5380.9434.411.04.24.232.036.0105.073.03.937.119.00.492.038.45.29.219.418.17.0Varuna × R1 B2-8186.36.87.449.0374.6423.611.83.74.135.039.0104.069.03.539.223.50.402.235.35.49.519.615.85.6Varuna × R1 B2-6201.07.27.550.5372.5423.011.93.54.133.037.0104.771.73.636.319.90.452.138.25.39.419.516.46.3Varuna × R1 B2-29195.78.37.453.0365.2418.212.83.64.231.735.7105.774.03.841.623.60.432.038.55.29.219.418.06.9Varuna × R1 B2-25204.09.47.453.3381.5434.811.94.34.432.336.3105.373.02.942.824.60.431.939.05.17.919.418.27.1R1 B2-26 × R1 B2-23203.28.310.454.5389.7444.212.84.84.331.335.3105.774.34.051.532.70.371.940.75.19.219.418.87.7R1 B2-26 × R1 B2-8200.34.95.549.5362.9412.411.44.64.335.041.0105.370.33.538.523.20.402.239.25.49.519.615.46.0R1 B2-26 × R1 B2-6201.68.69.453.5386.6440.113.14.64.332.038.0104.772.73.949.931.80.361.941.35.09.219.418.17.5R1 B2-26 × R1 B2-29192.78.88.353.5401.8455.311.84.94.331.737.7105.373.73.943.725.50.421.943.05.19.219.418.27.8R1 B2-26 × R1 B2-25192.99.711.454.8415.3470.112.54.44.431.337.3105.374.04.047.328.30.401.943.35.09.119.418.98.2T 6342 × R1 B2-23203.36.79.349.2359.7408.812.54.64.334.740.7104.770.03.452.036.40.302.237.35.49.519.615.65.8T 6342 × R1 B2-8180.26.88.446.5305.2351.711.63.84.336.342.3104.768.33.056.242.70.242.434.95.69.719.713.54.7T 6342 × R1 B2-6199.97.08.451.0357.8408.812.44.24.334.340.3105.771.33.648.832.50.332.038.35.39.419.516.36.3T 6342 × R1 B2-29202.29.58.753.1387.2440.312.25.24.332.338.3107.074.73.943.325.40.412.038.75.39.319.417.96.9T 6342 × R1 B2-25204.39.59.453.5386.2439.713.24.44.431.737.7106.374.73.045.627.10.411.839.15.19.219.418.57.2Punjab Local × R1 B2-23200.07.84.541.5272.1313.612.05.24.239.745.7104.765.02.434.524.30.302.537.85.710.020.010.33.9Punjab Local × R1 B2-8163.65.34.937.0189.1226.111.85.04.244.749.7106.762.01.721.214.30.332.834.65.810.420.36.92.4Punjab Local × R1 B2-6180.36.97.643.5293.3336.812.64.84.239.344.3107.067.72.632.721.40.352.637.95.79.919.911.34.3Punjab Local × R1 B2-29187.17.28.346.5311.5358.011.85.34.337.742.7107.069.32.950.337.50.252.338.35.49.819.712.84.9Punjab Local × R1 B2-25204.08.87.350.5356.9407.412.14.14.335.040.0105.370.32.545.529.60.352.238.85.49.419.515.96.2Mean (Hybrids)189.78.17.849.8357.5407.312.34.44.234.139.1105.471.23.344.228.10.372.138.65.39.419.516.16.2Grand mean185.37.77.448.5308.2356.712.44.44.234.038.9106.872.83.242.727.80.352.238.45.49.519.614.95.8SE (±)5.60.30.43.67.57.30.20.20.00.71.10.81.10.12.82.80.030.21.60.20.40.30.60.3CD (P=0.0511.10.60.87.115.014.60.50.50.11.32.11.62.10.35.65.60.060.33.10.40.70.71.20.6PH: plant height (cm), PRB: number of total primary branches per plant, SEB: number of total secondary branches per plant, SMR: number of siliquae on the main raceme, SB: number of siliquae on branches, TOS: total number of siliquae per plant, SDS: no. seeds per siliqua, SL: length of siliqua (cm), TW: test weight (g), DFIF: days to 1st flowering, DFF: days to 50% flowering, DM: days to maturity∗∗, MD: duration of maturity (i.e., duration from 1st flowering to maturity), YTT: yield of the top 10 cm of the main raceme (g), BIY: biological yield per plant (g), SY: stick yield per plant(g), HI: harvest index, AII: aphid infestation index, OIC: oil content (%), PHC: phenol content (mg/g), SUC: sugar content (mg/g), PRC: protein content (%), YPP: yield per plant (g), OYP: oil yield per plant (g).The analysis of variance disclosed that all features under examination had a significant genotype effect. This demonstrates the existence of adequate genetic diversity among lines, testers, and hybrids and enables a more thorough evaluation of the general combining ability studies (Tables3–5). Combining ability analysis is commonly used in self-pollinated crops to identify suitable parents and, more specifically, a cross with a higher possibility for producing desirable recombinant lines. The combining ability study demonstrated that the mean squares for all twenty-four yield and quality features were highly significant among parents, indicating the existence of genetic diversity between the parents (Table 2). The hybrids differed significantly for all parameters, demonstrating genetic diversity, except for days to maturity and protein content, which have modest variability. For all parameters except siliqua length and stick yield per plant, the variation due to parents versus hybrids was remarkable, showing the presence of significant variations between the crosses [36].Table 3
Analysis of variance for combining ability effects of yield and its attributed traits for parents and hybrids in mustard.
Source of variationdfPHPRBSEBSMRSBTOSSDSSLReplication2109.371NS0.334NS0.242NS6.133NS59.114NS56.604NS0.089NS0.091NSGenotypes46858.6776.553∗∗8.257∗∗94.504∗∗26824.021∗∗28792.752∗∗1.499∗∗0.794∗∗Parents111522.963∗∗3.258∗∗3.026∗∗138.196∗∗4967.440∗∗5230.693∗∗2.682∗∗1.220∗∗Parents vs crosses18065.407∗∗70.451∗∗54.977∗∗769.322∗∗997557.780∗∗1053732.616∗∗8.754∗∗0.094NSCrosses34431.798∗∗5.740∗∗8.576∗∗60.520∗∗5344.275∗∗6270.481∗∗0.903∗∗0.677∗∗Lines6805.998NS7.777∗14.400∗131.694∗∗17861.705∗∗20937.150∗∗0.995NS1.900∗∗Testers4524.105NS23.418∗∗20.480∗150.066∗∗10247.879∗∗12689.890∗∗0.827NS0.666NSLine × testers24322.864∗∗2.284∗∗5.136∗∗27.802NS1397.650∗∗1533.912∗∗0.892∗∗0.373∗∗Error9246.6310.1160.22119.11885.42480.6910.0910.091NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 4
Analysis of variance for combining ability effects of yield and its attributed traits for parents and hybrids in mustard.
Source of variationdfTWDFIFDFFDMMDYTTBIYSYReplication20.002NS0.178NS1.012NS1.114NS2.151NS0.004NS2.297NS3.827NSGenotypes460.084∗∗34.951∗∗37.461∗∗40.109∗∗71.908∗∗1.401289.527∗∗194.535∗∗Parents110.107∗∗68.677∗∗67.452∗∗84.613∗∗138.357∗∗2.373∗∗502.125∗∗295.775∗∗Parents vs crosses10.873∗∗4.177∗12.014∗∗860.857∗∗984.963∗∗8.398∗∗916.621∗∗30.804NSCrosses340.054∗∗24.945∗∗28.506∗∗1.571NS23.556∗∗0.881∗∗202.301∗∗166.596∗∗Lines60.105∗90.022∗∗104.200∗∗2.905NS70.327∗∗3.304∗∗524.668∗365.777∗Testers40.127∗∗55.938∗∗59.65∗∗71.929NS68.938∗∗1.593∗∗31.093NS52.209NSLinex testers240.029∗∗3.510∗∗4.390∗∗1.179NS4.299∗∗0.156∗∗150.244∗∗135.866∗∗Error920.0030.6911.7201.0091.7320.03411.89912.008NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 5
Analysis of variance for combining ability effects of yield and its attributed traits for parents and hybrids in mustard.
Source of variationdfHIAIIOICPHCSUCPRCYPPOYPReplication20.001NS0.037NS1.273NS0.010NS0.444NS0.014NS0.340NS0.019NSGenotypes460.014∗∗0.360∗∗11.824∗∗0.187∗∗0.511∗∗0.485∗∗38.431∗∗7.065∗∗Parents110.011∗∗0.959∗∗9.648∗∗0.404∗∗0.370NS1.529∗∗41.377∗∗6.748∗∗Parents vs crosses10.112∗∗1.132∗∗27.862∗∗0.313∗4.681∗∗1.540∗∗590.205∗∗99.128∗∗Crosses340.012∗∗0.143∗∗12.057∗∗0.114∗∗0.434∗∗0.116NS21.249∗∗4.460∗∗Lines60.031∗∗0.457∗∗30.566∗∗0.340∗∗1.084∗∗0.404∗∗73.904∗∗14.012∗∗Testers40.019∗0.38∗∗052.368∗∗0.281∗∗1.120∗∗0.224∗∗50.064∗∗14.486∗∗Linex testers240.007∗∗0.026NS0.711NS0.029NS0.157NS0.027NS3.283∗∗0.400∗∗Error920.0010.0393.6620.0540.2110.1710.5050.141NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Except for plant height, seed number per siliqua, and days to maturity, the lines showed significant differences in all studied characteristics, whereas the testers showed significant differences in the majority of the studied characteristics, except for plant height, seed number per siliqua, siliqua length, day to maturity, and biological and stick yield per plant. Except for plant height, seed number per siliqua, days to maturity, siliqua length, biological and stick yield per crop, additive variance predominated for the majority of the parameters [37].Due to the average squares, the line × tester interaction constituents were also remarkable for all yield components, specifying that merging ability played a significant role in the declaration of these traits and served as a direct test for the significance of dominant or nonadditive variance for the characters under study. However, the same was not found for certain less significant characters such as the total number of siliquae on the main raceme, days to maturity, aphid infestation index, oil content, and other biochemical components such as phenol, sugar, and protein.


### 3.2. General and Specific Combining Abilities

R1 B2-26 was the best combiner for oil yield/plant among the seven lines, although Varuna and JD6 both had significant GCA values. Punjab Local, on the other hand, exhibited a significantly negative GCA, while Kranti, R1 B2-5, and T 6342 had nonsignificant GCAs (Tables6 and 7). This means that by employing R1 B2-26, Varuna, and JD6 as parents, oil yield/plant may be increased; for instance, the number of primary and secondary branches on a plant, the number of siliquae on a raceme, the number of seeds on a raceme, the number of seeds per siliquae on a plant, the number of seeds per siliquae on a plant, the number of seeds per siliquae on a plant, the number of seeds per siliquae on a plant, and the number of seeds per siliquae on a plant. Additionally, R1 B2-26 demonstrated statistically significant GCA values for plant height, secondary branch count, siliqua count (main raceme, branches, and total), siliqua length, weight, maturity duration, biological yield (top 10 cm of the main raceme), harvest index (oil content), and seed production per plant.Table 6
General combining ability (GCA) effects of parents on yield and its attributed traits in mustard.
ParentsPHPRBSEBSMRSBTOSSDSSLTWDFIFDFFDMLines (gi)Kranti2.48NS1.46∗∗0.40∗∗0.40NS1.94NS2.34NS0.29∗∗0.19∗0.03NS−0.20NS−0.07NS0.15NSJD 6−3.84∗0.10NS0.09NS−0.11NS19.36∗∗19.26∗∗0.30∗∗−0.29∗∗0.01NS−2.00∗∗−2.27∗∗−0.11NSR1 B2-5−12.41∗∗−0.09NS−1.14∗∗−0.44NS−1.37NS−1.81NS−0.11NS−0.18∗−0.16∗∗0.53∗−0.40NS−0.51NSVaruna−0.32NS−0.23∗∗−0.36∗∗2.03NS17.45∗∗19.49∗∗−0.38∗∗−0.54−0.04∗∗−1.33∗∗−2.27∗∗−0.45NSR1 B2-268.46∗∗−0.08NS1.23∗∗3.33∗∗33.78∗∗37.11∗∗0.04NS0.29∗∗0.08∗∗−1.87∗∗−1.20∗∗−0.11NST 63428.29∗∗−0.23∗∗1.050.82NS1.73NS2.55NS0.09NS0.05NS0.08∗∗−0.27NS0.80∗0.29NSPunjab Local−2.68NS−0.92∗∗−1.25∗∗−6.04∗∗−72.91∗∗−78.94∗∗−0.23∗∗0.48∗∗0.01NS5.13∗∗5.40∗∗0.75∗∗SE (gi)1.760.090.121.132.392.320.080.080.010.210.340.26Tester (gj)R1 B2-23−0.80NS−0.35∗∗−0.28∗∗−2.15∗−3.47NS−5.62∗∗−0.16∗0.24∗∗−0.03∗∗0.10NS0.03NS−0.29NSR1 B2-8−6.56∗∗−1.49∗∗−1.57∗∗−3.12∗∗−34.22∗∗−37.35∗∗−0.24∗∗−0.20∗∗−0.09∗∗2.53∗∗2.60∗∗−0.24NSR1 B2-61.42NS−0.14NS0.63∗∗0.24NS2.09NS2.33NS0.15∗−0.09NS−0.02NS0.10NS0.22NS0.05NSR1 B2-29−1.31NS0.71∗∗0.29∗∗1.66NS9.72∗∗11.38∗∗0.03NS0.13∗0.02NS−0.90∗∗−0.92∗∗0.48∗R1 B2-257.24∗∗1.26∗∗0.93∗∗3.37∗∗25.88∗∗29.25∗∗0.22∗∗−0.08NS0.12∗∗−1.85∗∗−1.92∗∗0.00NSSE (gj)1.490.070.100.952.021.960.070.070.010.180.290.22NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 7
General combining ability (GCA) effects of parents on yield and its attributed traits in mustard.
ParentsMDYTTBIYSYHIAIIOICPHCSUCPRCYPPOYPLines (gi)Kranti0.35NS0.02NS5.67∗∗5.54∗∗−0.03∗∗0.01NS−0.68NS0.02NS0.06NS−0.03NS0.13NS−0.08NSJD 61.89∗∗0.32∗∗5.71∗∗3.99∗∗−0.01NS−0.12∗−0.12NS−0.08NS−0.17NS−0.12NS1.72∗∗0.62∗∗R1 B2-5−1.05∗∗−0.12∗−6.30∗∗−5.86∗∗0.04∗∗0.05NS0.82NS0.07NS0.08NS−0.01NS−0.44∗−0.06NSVaruna0.89∗0.20∗∗−4.77∗∗−5.97∗∗0.07∗∗−0.09NS−0.77NS−0.09NS−0.34∗∗−0.08NS1.20∗∗0.31∗∗R1 B2-261.75∗∗0.51∗∗2.01∗0.22NS0.02∗−0.18∗∗2.87∗∗−0.20∗∗−0.15NS−0.11NS1.79∗∗1.19∗∗T 63420.550.03NS5.00∗∗4.74∗∗−0.03∗∗−0.03NS−0.96NS−0.01NS0.02NS−0.01NS0.26NS−0.06NSPunjab Local−4.38∗∗−0.95∗∗−7.32∗∗−2.66∗∗−0.06∗∗0.35∗∗−1.15∗0.28∗∗0.51∗∗0.36−4.66∗∗−1.92∗∗SE (gi)0.340.050.890.890.010.050.490.060.120.110.180.10Tester (gj)R1 B2-23−0.39NS0.13∗∗0.51NS0.64NS−0.01NS0.01NS−0.36NS0.01NS0.04NS0.01NS−0.13NS−0.13NSR1 B2-8−2.77∗∗−0.27∗∗−0.97NS1.32NS−0.04∗∗0.20∗∗−2.50∗∗0.18∗∗0.27∗∗0.15NS−2.29∗∗−1.24∗∗R1 B2-6−0.06NS0.14∗∗−0.90NS−0.67NS0.00NS0.02NS0.30NS0.01NS0.09NS0.01NS−0.23NS−0.06NSR1 B2-291.37∗∗0.31∗∗1.90∗1.16NS0.00NS−0.06NS0.97∗−0.06NS−0.04NS−0.05NS0.75∗∗0.43∗∗R1 B2-251.85∗∗−0.31∗∗−0.54NS−2.45∗∗0.04∗∗−0.17∗∗1.60∗∗−0.14∗∗−0.36∗∗−0.13NS1.91∗∗1.00∗∗SE (gi)0.290.040.750.760.010.040.420.050.100.090.160.08NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.For JD6 and Varuna, GCA values for maturity, the top 10 cm of the main raceme yield, and yield per plant were all considerably positive. However, both had a negative impact of GCA on siliqua length, first flowering day, and 50% flowering days. By reducing the height of the plant and aphid infection index, JD6 caused dwarfism and aphid resistance. For days to maturity, both JD6 and Varuna showed high GCA values, indicating an unfavorable increase in crop length. Only Punjab Local had the required significantly negative GCA value for oil yield per plant and plant height. Utilizing this line to enhance the character should be done with caution since it has the potential to significantly reduce the yield.R1 B2-25 and R1 B2-29 had significant positive GCA values for oil yield/plant, whereas R1 B2-8 had significant negative GCA values. When R1 B2-25 and R1 B2-29 are used as parents, the character is enhanced; however, when R1 B2-8 is used as a parent, the oil yield/plant is decreased. Other yield-related characteristics such as plant height, number of primary and secondary branches per plant, number of siliquae on the main raceme, number of siliquae on branches, number of seeds per siliqua, weight, maturity duration, harvest index, oil content, and seed yield per plant all showed significant positive GCA values. R1 B2-29 had a significant positive correlation between the total number of primary and secondary branches on a plant, siliquae on branches, total siliqua per plant, length of siliqua, days to maturity, maturity duration, yield of the top 10 cm of the main raceme, biological yield per plant, oil content, and seed yield per plant. R1 B2-8 has a substantial negative GCA when compared with plant height. The number of major and secondary branches, the number of siliquae on the main raceme and on branches, the number of siliquae per plant, the number of seeds per siliqua, the length of siliqua (cm), and the weight of siliqua are all factors to consider. Despite the presence of dwarfing genes in R1 B2-8, utilizing it to improve features should be done with caution since adopting this line may eventually result in a considerable drop in yield and other yield-related qualities like aphid resistance.However, in JD6, R1 B2-26 between the lines can be chosen as an expected parent for increasing oil yield/plant, secondary branch number per plant, siliqua on the main raceme, siliqua on branches, total siliqua per plant, siliqua length, weight, top 10 cm of the main raceme yield, biological yield per crop, and harvest.R1 B2-25 may be considered as a possible parent to enhance oil yield, aphid resistance, and seed size among the testers, as evidenced by a significant positive GCA for test weight, although it may delay maturity. This parent, on the other hand, may boost the amount of oil, phenol, and sugar. Despite having a strong GCA influence on oil content, R1 B2-29 is a poor general combiner.While SCA effects do not directly benefit self-pollinated crops, ideal hybrids should provide transgressive segregants that can be selected as potent homozygous lines.Punjab Local × R1 B2-25 and Varuna × R1 B2-23 were the only crossings with significantly positive SCA values for oil yield/plant (Tables8–12). On the other hand, three crossings (Punjab Local × R1 B2-8, Varuna × R1 B2-25, and JD6 × R1 B2-25) had considerably negative SCA for plant yield. Plant height, number of siliquae on branches, number of siliquae per plant, maturity, yield of the top 10 cm of the main raceme, biological yield per plant, stick yield per plant, and seed yield per plant have all exhibited a significant positive SCA effect in the cross Punjab Local × R1 B2-8. This hybrid, on the other hand, showed significant SCA impacts on plant height and seed yield per siliqua.Table 8
Specific combining ability (SCA) effects of mustard hybrids on yield and its attributed traits.
HybridsPHPRBSEBSMRSBKranti × R1 B2-23−6.704NS0.717∗∗−0.344NS2.419NS−3.344NSKranti × R1 B2-84.391NS1.389∗∗0.587∗0.721NS23.478∗∗Kranti × R1 B2-66.410NS0.070NS1.382∗∗−0.969NS−6.575NSKranti × R1 B2-29−2.390NS−0.778∗∗−1.208∗∗−1.731NS−8.132NSKranti × R1 B2-25−1.709NS−1.397∗∗−0.418NS−0.440NS−5.427NSJD 6 × R1 B2-234.950NS−1.556∗∗−0.897∗∗−10.461∗∗13.183∗JD 6 × R1 B2-86.045NS−0.318NS−2.600∗∗4.058NS13.521∗JD 6 × R1 B2-6−2.303NS1.163∗∗0.696∗3.534NS−2.265NSJD 6 × R1 B2-29−9.603∗0.615∗∗1.472∗∗2.339NS−7.355NSJD 6 × R1 B2-250.911NS0.096NS1.329∗∗0.530NS−17.084∗∗R1 B2-5 × R1 B2-234.756NS−0.103NS0.243NS0.252NS−16.097∗∗R1 B2-5 × R1 B2-85.451NS0.335NS1.324∗∗0.221NS4.358NSR1 B2-5 × R1 B2-6−8.696∗−0.384−0.681∗−0.636NS17.039∗∗R1 B2-5 × R1 B2-291.504NS0.169NS0.062NS−0.564NS−1.419NSR1 B2-5 × R1 B2-25−3.015NS−0.017NS−0.9480.726NS−3.880NSVaruna × R1 B2-23−28.704∗∗0.210NS0.316NS3.786NS9.443NSVaruna × R1 B2-83.525NS0.415∗1.514∗∗0.255NS33.898∗∗Varuna × R1 B2-610.177∗−0.604∗∗−0.524NS−1.602NS−4.521NSVaruna × R1 B2-297.610NS−0.285NS−0.315NS−0.531NS−19.479∗∗Varuna × R1 B2-257.391NS0.263NS−0.991∗∗−1.907NS−19.340∗∗R1 B2-26 × R1 B2-235.823NS0.6241.663∗∗3.486NS1.883NSR1 B2-26 × R1 B2-88.685∗−1.705−1.906∗∗−0.545NS5.838NSR1 B2-26 × R1 B2-62.037NS0.643−0.244NS0.098NS−6.748NSR1 B2-26 × R1 B2-29−4.096NS0.062NS−1.001∗∗−1.331NS0.861NSR1 B2-26 × R1 B2-25−12.449∗∗0.376NS1.489∗∗−1.707NS−1.834NST 6342 × R1 B2-236.090NS−0.856∗∗0.743∗∗0.666NS3.930NST 6342 × R1 B2-8−11.249∗∗0.349NS1.107∗∗−1.032NS−19.816∗∗T 6342 × R1 B2-60.504NS−0.737∗∗−1.064∗∗0.111NS−3.535NST 6342 × R1 B2-295.537NS0.915∗∗−0.455NS0.782NS18.275∗∗T 6342 × R1 B2-25−0.882NS0.330NS−0.331NS−0.527NS1.146NSPunjab Local × R1 B2-2313.790∗∗0.964∗∗−1.724∗∗−0.148NS−8.997NSPunjab Local × R1 B2-8−16.849∗∗−0.465∗−0.026NS−3.679NS−61.276∗∗Punjab Local × R1 B2-6−8.130∗−0.150NS0.436NS−0.536NS6.605NSPunjab Local × R1 B2-291.437NS−0.698∗∗1.445∗∗1.036NS17.248∗∗Punjab Local × R1 B2-259.751∗0.350NS−0.131NS3.326NS46.420∗∗SE (±)3.9430.1970.2722.5245.336NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 9
Specific combining ability (SCA) effects of mustard hybrids on yield and its attributed traits.
HybridsTOSSDSSLTWDFIFKranti × R1 B2-23−0.925NS−0.625∗∗0.434∗0.123∗∗−0.371NSKranti × R1 B2-824.199∗∗0.429∗−0.414∗−0.115∗∗−0.800NSKranti × R1 B2-6−7.544NS−0.012NS0.376∗−0.025NS0.629NSKranti × R1 B2-29−9.863NS0.209NS−0.305NS0.049NS0.295NSKranti × R1 B2-25−5.868NS−0.001NS−0.090NS−0.031NS0.248NSJD 6 × R1 B2-232.722NS−0.106NS−0.429∗0.147∗∗0.095NSJD 6 × R1 B2-817.579∗∗−0.280NS0.247NS0.022NS−1.000∗JD 6 × R1 B2-61.270NS−0.522∗∗0.008NS−0.065∗−0.571NSJD 6 × R1 B2-29−5.016NS0.423∗−0.165NS−0.074∗0.429NSJD 6 × R1 B2-25−16.554∗∗0.484∗∗0.340NS−0.030NS1.048∗R1 B2-5 × R1 B2-23−15.845∗∗0.564∗∗0.047NS−0.315∗∗1.229∗R1 B2-5 × R1 B2-84.579NS0.859∗∗0.061NS−0.032NS−0.867NSR1 B2-5 × R1 B2-616.403∗∗−0.210NS−0.010NS0.091∗∗−0.105NSR1 B2-5 × R1 B2-29−1.983−0.529∗∗−0.1420.088∗∗0.562R1 B2-5 × R1 B2-25−3.154NS−0.684∗∗0.044NS0.168∗∗−0.819NSVaruna × R1 B2-2313.229∗−0.760∗∗0.072NS0.027NS−0.905NSVaruna × R1 B2-834.152∗∗0.175NS0.082NS0.002NS−0.333NSVaruna × R1 B2-6−6.124NS−0.121NS−0.246NS−0.048NS0.095NSVaruna × R1 B2-29−20.010∗∗0.887−0.414∗−0.034NS−0.238NSVaruna × R1 B2-25−21.248∗∗−0.1810.507∗∗0.053NS1.381∗∗R1 B2-26 × R1 B2-235.369NS0.606∗∗−0.079NS0.011NS−1.038∗R1 B2-26 × R1 B2-85.292NS−0.680∗∗0.133NS0.043NS0.200NSR1 B2-26 × R1 B2-6−6.650NS0.605∗∗0.004NS0.019NS−0.371NSR1 B2-26 × R1 B2-29−0.470NS−0.507∗∗0.095NS−0.013NS0.295NSR1 B2-26 × R1 B2-25−3.541NS−0.024NS−0.153NS−0.060NS0.914NST 6342 × R1 B2-234.595NS0.244NS−0.120NS0.015NS0.695NST 6342 × R1 B2-8−20.848∗∗−0.495∗∗−0.473∗∗0.048NS−0.067NST 6342 × R1 B2-6−3.424NS−0.123NS−0.134NS0.011NS0.362NST 6342 × R1 B2-2919.057∗∗−0.235NS0.674−0.008NS−0.638NST 6342 × R1 B2-250.619NS0.609∗∗0.053NS−0.065∗−0.352NSPunjab Local × R1 B2-23−9.145NS0.077NS0.075NS−0.007NS0.295NSPunjab Local × R1 B2-8−64.954∗∗−0.008NS0.364∗0.032NS2.867∗∗Punjab Local × R1 B2-66.070NS0.383∗0.003NS0.018NS−0.038NSPunjab Local × R1 B2-2918.284∗∗−0.249NS0.257NS−0.008NS−0.705NSPunjab Local × R1 B2-2549.746∗∗−0.204NS−0.700∗∗−0.034NS−2.419∗∗SE (±)5.1860.1740.1740.0310.480NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 10
Specific combining ability (SCA) effects of mustard hybrids on yield and its attributed traits.
HybridsDFFDMMDYTTBIYKranti × R1 B2-23−0.362NS0.086NS0.457NS0.078NS−4.247∗Kranti × R1 B2-8−0.933NS0.705NS1.505NS0.200NS11.362∗∗Kranti × R1 B2-60.448NS0.086NS−0.543NS−0.052NS8.124∗∗Kranti × R1 B2-290.590NS−0.676NS−0.971NS−0.103NS−5.642∗∗Kranti × R1 B2-250.257NS−0.200NS−0.448NS−0.122NS−9.597∗∗JD 6 × R1 B2-230.505NS0.352NS0.257NS0.107NS2.279NSJD 6 × R1 B2-8−0.733NS−0.029NS0.971NS0.152NS1.801NSJD 6 × R1 B2-6−0.019NS0.019NS0.590NS0.110NS−7.650∗∗JD 6 × R1 B2-29−0.210NS−0.076NS−0.505NS0.013NS7.134∗∗JD 6 × R1 B2-250.457NS−0.267NS−1.314NS−0.383∗∗−3.564NSR1 B2-5 × R1 B2-231.305NS0.419NS−0.810NS−0.182NS0.463NSR1 B2-5 × R1 B2-8−0.933NS−0.295NS0.571NS0.123NS−0.598NSR1 B2-5 × R1 B2-6−0.219NS0.086NS0.190NS0.021NS−0.206NSR1 B2-5 × R1 B2-290.590NS−0.343NS−0.905NS0.017NS−1.112NSR1 B2-5 × R1 B2-25−0.743NS0.133NS0.952NS0.021NS1.453NSVaruna × R1 B2-23−0.829NS0.352NS1.257NS0.256∗−2.790NSVaruna × R1 B2-8−0.400NS−0.695NS−0.362NS0.185NS0.799NSVaruna × R1 B2-6−0.019NS−0.314NS−0.410NS−0.087NS−2.199NSVaruna × R1 B2-29−0.210NS0.257NS0.495NS−0.061NS0.275NSVaruna × R1 B2-251.457NS0.400NS−0.981NS−0.293∗∗3.916NSR1 B2-26 × R1 B2-23−2.562∗∗0.686NS1.724∗−0.024NS4.816∗R1 B2-26 × R1 B2-80.533NS0.305NS0.105NS−0.106NS−6.661∗∗R1 B2-26 × R1 B2-6−0.086NS−0.648NS−0.276NS−0.081NS4.608∗R1 B2-26 × R1 B2-290.724NS−0.410NS−0.705NS−0.222∗−4.402∗R1 B2-26 × R1 B2-251.390NS0.067−0.848NS0.433∗∗1.640NST 6342 × R1 B2-230.771NS−0.714−1.410NS−0.071NS2.287NST 6342 × R1 B2-8−0.133NS−0.762NS−0.695NS−0.116NS7.976∗∗T 6342 × R1 B2-60.248NS−0.048NS−0.410NS0.062NS0.575NST 6342 × R1 B2-29−0.610NS0.857NS1.495NS0.195NS−7.791∗∗T 6342 × R1 B2-25−0.276NS0.667NS1.019NS−0.071NS−3.046NSPunjab Local × R1 B2-231.171NS−1.181∗−1.476NS−0.163NS−2.807NSPunjab Local × R1 B2-82.600∗∗0.771NS−2.095∗∗−0.438∗∗−14.678∗∗Punjab Local × R1 B2-6−0.352NS0.819NS0.857NS0.027NS−3.252NSPunjab Local × R1 B2-29−0.876NS0.390NS1.095NS0.160NS11.538∗∗Punjab Local × R1 B2-25−2.543∗∗−0.800NS1.619∗0.414∗∗9.200∗∗SE (±)0.7570.5800.7600.1061.992NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 11
Specific combining ability (SCA) effects of mustard hybrids on yield and its attributed traits.
HybridsSYHIAIIOICPHCKranti × R1 B2-23−4.552∗0.029NS−0.052NS−1.016NS−0.045NSKranti × R1 B2-810.433∗∗−0.050∗−0.038NS0.089NS−0.018NSKranti × R1 B2-68.586∗∗−0.068∗∗0.038NS0.288NS0.049NSKranti × R1 B2-29−4.971∗0.016NS0.057NS0.017NS0.050NSKranti × R1 B2-25−9.495∗∗0.074∗∗−0.005NS0.622NS−0.037NSJD 6 × R1 B2-231.934NS−0.010NS−0.052NS−0.483NS−0.077NSJD 6 × R1 B2-81.087NS−0.003NS−0.138NS−0.111NS−0.150NSJD 6 × R1 B2-6−8.094∗∗0.070∗∗−0.029NS−0.145NS−0.050NSJD 6 × R1 B2-297.249∗∗−0.051∗0.090NS0.484NS0.051NSJD 6 × R1 B2-25−2.175NS−0.006NS0.129NS0.255NS0.225NSR1 B2-5 × R1 B2-231.488NS−0.030NS0.114NS0.417NS0.108NSR1 B2-5 × R1 B2-8−1.260NS0.013NS−0.071NS0.455NS−0.064NSR1 B2-5 × R1 B2-6−0.141NS0.001NS0.005NS−0.078NS0.002NSR1 B2-5 × R1 B2-29−0.965NS0.006NS−0.010NS−0.583NS−0.030NSR1 B2-5 × R1 B2-250.878NS0.010NS−0.038NS−0.211NS−0.017NSVaruna × R1 B2-23−3.772NS0.059∗∗−0.052NS0.904NS−0.076NSVaruna × R1 B2-80.047NS0.001NS−0.038NS−0.058NS−0.008NSVaruna × R1 B2-6−1.568NS0.009NS0.038NS0.008NS0.058NSVaruna × R1 B2-290.342NS−0.009NS0.024NS−0.363NS0.019NSVaruna × R1 B2-254.951∗−0.059∗∗0.029NS−0.491NS0.006NSR1 B2-26 × R1 B2-233.734NS−0.013NS−0.092NS−0.469NS−0.061NSR1 B2-26 × R1 B2-8−6.447∗∗0.049∗0.055NS0.202NS0.106NSR1 B2-26 × R1 B2-64.206∗−0.031NS−0.069NS−0.465NS−0.100NSR1 B2-26 × R1 B2-29−3.985∗0.027NS0.017NS0.497NS0.041NSR1 B2-26 × R1 B2-252.491NS−0.032NS0.089NS0.235NS0.014NST 6342 × R1 B2-232.948NS−0.029NS0.094NS−0.036NS0.090NST 6342 × R1 B2-88.567∗∗−0.060∗∗0.109NS−0.231NS0.117NST 6342 × R1 B2-60.352NS−0.008NS−0.082NS0.302NS−0.083NST 6342 × R1 B2-29−8.605∗∗0.074∗∗−0.030NS0.097NS0.011NST 6342 × R1 B2-25−3.262NS0.023NS−0.091NS−0.131NS−0.135NSPunjab Local × R1 B2-23−1.779NS−0.007NS0.041NS0.683NS0.062NSPunjab Local × R1 B2-8−12.427∗∗0.050∗0.122NS−0.345NS0.016NSPunjab Local × R1 B2-6−3.341NS0.027NS0.098NS0.091NS0.122NSPunjab Local × R1 B2-2910.935∗∗−0.062∗∗−0.150NS−0.150NS−0.143NSPunjab Local × R1 B2-256.611∗∗−0.009NS−0.111NS−0.279NS−0.057NSSE (±)2.0010.0200.1141.1050.135NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 12
Specific combining ability (SCA) effects of mustard hybrids on yield and its attributed traits.
HybridsSUCPRCYPPOYPKranti × R1 B2-23−0.079NS−0.013NS0.305NS−0.043NSKranti × R1 B2-8−0.162NS−0.087NS0.929∗0.353NSKranti × R1 B2-60.131NS0.011NS−0.461NS−0.125NSKranti × R1 B2-29−0.005NS0.055NS−0.670NS−0.256NSKranti × R1 B2-250.115NS0.034NS−0.102NS0.070NSJD 6 × R1 B2-23−0.058NS−0.022NS0.345NS0.048NSJD 6 × R1 B2-8−0.084NS−0.063NS0.715NS0.199NSJD 6 × R1 B2-6−0.114NS−0.022NS0.445NS0.159NSJD 6 × R1 B2-29−0.027NS0.025NS−0.114NS0.066NSJD 6 × R1 B2-250.283NS0.082NS−1.389∗∗−0.472∗R1 B2-5 × R1 B2-230.040NS0.034NS−1.025∗−0.331NSR1 B2-5 × R1 B2-8−0.110NS−0.083NS0.662NS0.299NSR1 B2-5 × R1 B2-6−0.040NS0.018NS−0.065NS−0.036NSR1 B2-5 × R1 B2-29−0.002NS0.028NS−0.147NS−0.147NSR1 B2-5 × R1 B2-250.111NS0.004NS0.575NS0.215NSVaruna × R1 B2-230.119NS−0.064NS0.982∗0.540∗Varuna × R1 B2-80.189NS−0.055NS0.752NS0.245NSVaruna × R1 B2-60.282NS0.043NS−0.631NS−0.227NSVaruna × R1 B2-290.230NS0.010NS−0.067NS−0.078NSVaruna × R1 B2-25−0.820∗∗0.066NS−1.035∗−0.481∗R1 B2-26 × R1 B2-23−0.105NS−0.052NS1.082∗∗0.353NSR1 B2-26 × R1 B2-8−0.055NS−0.016NS−0.215NS−0.165NSR1 B2-26 × R1 B2-6−0.105NS−0.032NS0.402NS0.087NSR1 B2-26 × R1 B2-29−0.001NS0.035NS−0.417NS−0.044NSR1 B2-26 × R1 B2-250.266NS0.065NS−0.852∗−0.231NST 6342 × R1 B2-230.016NS0.028NS−0.661NS−0.256NST 6342 × R1 B2-80.026NS0.034NS−0.591NS−0.237NST 6342 × R1 B2-6−0.081NS−0.015NS0.223NS0.130NST 6342 × R1 B2-29−0.117NS−0.038NS0.814NS0.320NST 6342 × R1 B2-250.156NS−0.008NS0.215NS0.044NSPunjab Local × R1 B2-230.066NS0.090NS−1.028∗−0.311NSPunjab Local × R1 B2-80.196NS0.270NS−2.251∗∗−0.695∗∗Punjab Local × R1 B2-6−0.074NS−0.002NS0.089NS0.012NSPunjab Local × R1 B2-29−0.077NS−0.115NS0.603NS0.139NSPunjab Local × R1 B2-25−0.110NS−0.242NS2.588∗∗0.855∗∗SE (±)0.2650.2390.4100.216NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Punjab Local × R1 B2-25 exhibited a strong unfavorable SCA impact at maturity. This cross is ideal for picking out the early offspring. Cross-combination with high means, correct SCA calculations, and at least one parent with a high GCA, according to Kenga et al. [38], would almost surely improve the desired trait concentration. In this study, Varuna showed a statistically insignificant negative GCA influence on oil yield per crop, but R1 B2-23 had a significant positive GCA effect.The parent Punjab Local had a detrimental effect on plant oil production via GCA. Additional significant GCA effects included siliqua length of the first flowering day to 50% flowering day and maturity days, aphid infestation index, phenol sugar, and protein content, as well as total per plant, primary and secondary branches, siliqua on the main raceme, siliqua on branches, total siliqua per plant, seed number per siliqua, and duration. In this study, significant negative GCA effects on all parameters except maturity time are considered undesirable. GCA has a negative impact on the first flowering day, 50% blooming day, maturity day, aphid infestation index, sugar and protein content, and aphid infestation index. Plant height, number of primary and secondary branches, total siliqua on the main raceme, total siliqua on branches, total number of siliquae per plant, number of seeds per siliqua, weight, maturity duration, harvest index, oil content, and seed yield per plant were all significantly improved by R1 B2-25.Varuna had significant positive GCA effects on oil yield per plant [39], siliqua yield per plant, yield of the top 10 cm of the main raceme, harvest index, and seed yield per crop, whereas Varuna had significant negative GCA effects. These characteristics benefit the breeder. It did, however, exhibit negative GCA effects on undesirable characteristics such as primary and secondary branching, seed number per siliqua, test weight, days to first and 50% flowering, and biological output per plant. The other parent of the cross, R1 B2-23, demonstrated no significant negative GCA effect on oil yield per crop, but a significant negative GCA effect on primary branches, secondary branches, siliqua on main strobili, total number of siliquae, seed number per siliqua, and test weight. As a result, one of the parents in this cross (Varuna) had a high GCA.The Varuna × R1 B2-23 hybrid has a negative effect on plant height because of its SCA content. This cross is perfect for reducing the height of plants. However, for a larger plant, Punjab Local × R1 B2-25 is preferable. Tiwari et al. [40] said that hybrid combinations including at least one good general combiner parent might be used to generate high-producing pure lines via additive gene action, even if SCA effects were negligible.In the present study, the variety Varuna exhibited a negative nonsignificant GCA effect for oil content. It also expressed a negative but nonsignificant general correlating ability effects for earliness and dwarfness, which is in contrary to the earlier findings [39]. This deviation in results may be due to the use of different breeding materials in different environments.Among the biochemical features, no cross exhibited a significant SCA value. Additionally, the aphid infestation index and oil content were similar. Only two hybrids showed a substantial SCA effect on oil production, showing that even if the parents’ grain yields varied widely, they produced hybrids with average oil yields higher than the parents’ limits, although little improvement.


### 3.3. Gene Action, Degree of Dominance, Heritability, Contribution to the Total Variance, and Heterosis

The level of variance attributable to line × testers was greater than that due to testers for the number of seeds per siliqua, biological yield per plant, and stick yield per plant, demonstrating nonadditive variance governing by these traits. To establish the relevance of the GCA and SCA variances produced from the line × tester design, these data appeared exploratory. Therefore, we can calculate the additive (σ2A) and nonadditive (σ2D) components of population variance and the predictability ratio (σ2A/(σ2A+σ2D)). The σ2gca/σ2sca ratio was less than one for all characters, indicating a major involvement of nonadditive gene actions such as dominance, epistasis, and other interaction effects in the expression of these characteristics. Mannuel and Palamisamy [41] and Bobby and Nadarajan [42] found similar results. Because the variance owing to general combining ability (σ2gca) was less than variance due to specific combining ability (σ2sca) for all traits investigated, this supports a predominance of nonadditive gene activity influencing these qualities (Tables 13 and 14). The variation ratio of general to specific combining ability (σ2gca/σ2sca) was less than unity, corroborating these findings. As a result, nonadditive gene results appeared to regulate the heritage of all studied features. Because recombinants within segregating populations might increase features including oil yield, plant height, number of siliquae per plant, and crop length, the selection of superior plants should be deferred to the following generations. In an earlier study using a variety of experimental lines, it was observed that the variance of GCA was greater in plant height and 1000-seed weight from sowing to 50% flowering and maturity, but the variation of SCA was more in seed yield and other remaining boundaries [43]. The discrepancy in the results is primarily attributable to the use of various testing materials. The dominance ratio suggested a predominance of nonadditive gene action in aphid resistance inheritance [44–50]. Negative variance was reported for (σ2D) aphid infestation index, oil content, phenol content, sugar content, and protein content, except for aphid infestation index. Given that they are squared values, the variance cannot be negative. However, they do occur, especially when estimating genetic variance using ANOVA expectation, as in this study to derive σ2gca from ANOVA for combining ability. There are a number of possible explanations for negative variance estimates: (a) small sample size and outlier values; (b) G-x-E interaction that may have increased the error variations; and (c) lack of randomly mating when producing half-sibs. Its validity can be accessed through a series of experiments using an average [51].Table 13
Estimates of genetic variance for yield and its attributed traits in mustard.
ParameterPHPRBSEBSMRSBTOSSDSSLTWDFIFDFFDMMDCovHS (Line)32.2090.3660.6186.9261097.6041293.5490.0070.1020.0055.7676.6540.1154.402CovHS (Tester)9.5831.0060.7315.822421.439531.237−0.0030.0140.0052.4972.6320.0363.078CovHS (Average)2.1290.0680.0670.63977.11892.5540.0000.0060.0000.4190.4710.0080.376CovFS163.8623.5464.23826.7453095.8713694.7680.2710.2840.02715.54017.1790.31614.622Ϭ2gca=2.1290.0680.0670.63977.11892.5540.0000.0060.0000.4190.4710.0080.376Ϭ2sca=92.0780.7231.6382.895437.409484.4070.2670.0940.0090.9400.8900.0560.856Ϭ2gca/Ϭ2sca0.0230.0930.0410.2210.1760.1910.0010.0630.0560.4460.5290.1360.440Ϭ2A=4.2570.1350.1341.279154.236185.1070.0000.0120.0010.8380.9420.0150.753Ϭ2D=92.0780.7231.6382.895437.409484.4070.2670.0940.0090.9400.8900.0560.856Ϭ2A/Ϭ2D=0.0460.1870.0820.4420.3530.3820.0020.1260.1120.8911.0590.2720.880Ϭ2A/(Ϭ2A+Ϭ2D)=0.0440.1570.0760.3060.2610.2760.0020.1120.1010.4710.5140.2140.468h2ns=0.0130.0600.0460.0290.0170.0190.0010.0370.0330.0690.0690.0010.030h2ns (%)=1.3425.9714.6362.8901.7141.9180.0733.6533.2516.9176.9130.1092.995Table 14
Estimates of genetic variance for yield and its attributed traits in mustard.
ParameterYTTBIYSYHIAIIOICPHCSUCPRCYPPOYPCovHS (Line)0.21024.96215.3270.0020.0291.9900.0210.0620.0254.7080.907CovHS (Tester)0.068−5.674−3.9840.0010.0172.4600.0120.0460.0092.2280.671CovHS (Average)0.0141.0170.6000.0000.0020.2220.0020.0050.0020.3510.079CovFS0.52272.44456.3360.0060.0787.6300.0510.1810.01213.2693.005Ϭ2gca=0.0141.0170.6000.0000.0020.2220.0020.0050.0020.3510.079Ϭ2sca=0.04146.11541.2860.002−0.004−0.984−0.008−0.018−0.0480.9260.087Ϭ2gca/Ϭ2sca0.3460.0220.0150.063−0.527−0.225−0.199−0.300−0.0360.3790.916Ϭ2A=0.0282.0341.2010.0000.0050.4430.0030.0110.0040.7020.159Ϭ2D=0.04146.11541.2860.002−0.004−0.984−0.008−0.018−0.0480.9260.087Ϭ2A/Ϭ2D=0.6920.0440.0290.126−1.055−0.451−0.397−0.600−0.0730.7581.831Ϭ2A/(Ϭ2A+Ϭ2D)=0.4090.0420.0280.11219.280−0.821−0.658−1.503−0.0790.4310.647h2ns=0.0580.0190.0160.0410.0320.0690.0330.0350.0130.0530.065h2ns (%)=5.7861.9481.6494.0973.1576.9473.3393.4771.2725.3416.478Furthermore, a predictability ratio greater than 0.5 denotes additive gene activation, whereas one less than 0.5 denotes nonadditive gene measurements. There are a lot of additive and nonadditive gene measurements in a characteristic. Predictability ratios were low and did not surpass 0.5 except for days to 50% flowering, aphid infestation index, and oil production per plant, showing nonadditive gene activity predominates. The prediction ratios for aphid infestation index and oil yield were higher, indicating the dominance of additive gene measures for these variables. This suggests that selection among segregating recombinants can result in plants that are superior in quality in regard to days to 50% blooming, aphid infestation index, and oil yield. All attributes had poor narrow-sense heritability values, according to the present study (Tables13 and 14).Line × testers donated more to the total sum square than any other factors for plant height, secondary branch count, seed count, days to maturity, biological yield, and stick yield (Table15 and Figure 1). Aphid infestation index, phenol and sugar content, seed yield per crop, and oil yield per crop were all determined by lines. In the presence of testers, the variation in primary branch count and oil content was more noticeable. In no case did all three sources of variance contribute equally. Testers and lines together contributed much less to plant height, seed number per siliqua, days to maturity, biological yield, and stick yield per crop. In addition, the interaction between lines and testers influenced the expression of the traits assessed. The current investigation found substantial differences in twenty-four traits between parents and hybrids (Figure 1). The contributions of lines and testers reflect a fixed component of genetic variation, whereas the line × the tester’s element reflects nonfixable factors [52].Table 15
Proportional contribution of lines, testers, and their interactions to total variance.
CharactersProportional contribution of lines, testers, and their interactions to total variance (in %)LinesTestersLines × testersPlant height (cm)32.9414.2852.78No. of primary branches per plant23.9148.0028.09No. of secondary branches per plant29.6328.1042.27No. of siliquae on main raceme38.4029.1732.43No. of siliquae on branches58.9822.5618.46No. of siliquae per plant58.9223.8117.27No. of seeds per siliqua19.4510.7869.77Length of siliqua (cm)49.5311.5738.90Test weight (g)34.3727.7237.92Days to 1st flowering63.6826.389.93Days to 50% flowering64.5124.6210.87Days to maturity32.6214.4452.94Duration of maturity52.6934.4312.88Yield of top 10 cm of the main raceme (g)66.2021.2812.52Biological yield per plant (g)45.771.8152.42Stick yield per plant (g)38.753.6957.57Harvest index44.2418.1537.61Aphid infestation index56.2031.1812.62Oil content (%)44.7451.104.16Phenol content (mg/g)52.6529.0718.28Sugar content (mg/g)44.0930.3525.57Protein content (%)61.2322.6116.16Yield per plant (g)61.3827.7210.91Oil yield per plant(g)55.4438.226.34Figure 1
Proportional contribution of lines, testers, and their interactions to total variance.A remarkable standard heterosis for oil yield per crop was observed in seven hybrids using JD6 as a commercial check: R1 B2-5 × R1 B2-25, Varuna × R1 B2-25, R1 B2-26 × R1 B2-6, and R1 B2-26 × R1 B2-26 (Tables16–20).Table 16
Standard (economic) heterosis of mustard hybrids for yield and its attributed traits.
CrossesPHPRBSEBSMRSBKranti × R1 B2-2315.00∗∗74.31∗∗37.96∗∗−1.22NS88.16∗∗Kranti × R1 B2-818.32∗∗66.15∗∗31.28∗∗−6.44NS86.06∗∗Kranti × R1 B2-624.55∗∗66.73∗∗85.97∗∗−3.18NS89.41∗∗Kranti × R1 B2-2917.36∗∗66.73∗∗32.49∗∗−1.87NS92.64∗∗Kranti × R1 B2-2523.11∗∗65.56∗∗58.62∗∗4.00NS102.71∗∗JD 6 × R1 B2-2318.32∗∗10.76∗22.16∗∗−27.40∗∗106.28∗∗JD 6 × R1 B2-815.41∗∗12.51∗−32.54∗∗−0.89NS90.05∗∗JD 6 × R1 B2-615.18∗∗62.07∗∗67.74∗∗4.65NS101.01∗∗JD 6 × R1 B2-298.94∗67.31∗∗75.64∗∗5.11NS102.36∗∗JD 6 × R1 B2-2520.81∗∗67.90∗∗84.76∗∗4.91NS105.79∗∗R1 B2-5 × R1 B2-2312.86∗∗32.92∗∗20.64∗∗−7.09NS79.59∗∗R1 B2-5 × R1 B2-89.70∗∗20.68∗∗16.69∗−9.04NS74.09∗∗R1 B2-5 × R1 B2-65.86NS31.75∗∗20.34∗∗−4.15NS100.24∗∗R1 B2-5 × R1 B2-2910.51∗∗56.24∗∗27.63∗∗−1.22NS94.46∗∗R1 B2-5 × R1 B2-2513.03∗∗62.65∗∗20.94∗∗4.65NS101.77∗∗Varuna × R1 B2-23−0.45NS35.83∗∗36.14∗∗4.65NS103.26∗∗Varuna × R1 B2-816.04∗∗19.51∗∗34.31∗∗−4.15NS99.90∗∗Varuna × R1 B2-625.15∗∗25.34∗∗37.35∗∗−1.22NS98.78∗∗Varuna × R1 B2-2921.85∗∗45.74∗∗34.92∗∗3.67NS94.87∗∗Varuna × R1 B2-2527.04∗∗64.98∗∗34.31∗∗4.32NS103.57∗∗R1 B2-26 × R1 B2-2326.52∗∗45.74∗∗89.62∗∗6.60NS107.94∗∗R1 B2-26 × R1 B2-824.71∗∗−14.89∗∗0.89NS−3.18NS93.64∗∗R1 B2-26 × R1 B2-625.54∗∗49.82∗∗71.39∗∗4.65NS106.31∗∗R1 B2-26 × R1 B2-2920.02∗∗54.49∗∗51.33∗∗4.65NS114.44∗∗R1 B2-26 × R1 B2-2520.15∗∗69.64∗∗108.46∗∗7.26NS121.62∗∗T 6342 × R1 B2-2326.58∗∗17.18∗∗69.56∗∗−3.83NS91.93∗∗T 6342 × R1 B2-812.20∗∗18.34∗∗52.55∗∗−9.04NS62.85∗∗T 6342 × R1 B2-624.48∗∗23.01∗∗53.15∗∗−0.24NS90.92∗∗T 6342 × R1 B2-2925.92∗∗66.73∗∗58.02∗∗3.87NS106.63∗∗T 6342 × R1 B2-2527.24∗∗66.15∗∗71.99∗∗4.65NS106.11∗∗Punjab Local × R1 B2-2324.55∗∗37.00∗∗−17.35∗−18.82∗∗45.20∗∗Punjab Local × R1 B2-81.88NS−7.89NS−10.05NS−27.63∗∗0.89NSPunjab Local × R1 B2-612.28∗∗21.26∗∗38.57∗∗−14.91∗56.50∗∗Punjab Local × R1 B2-2916.53∗∗26.50∗∗50.72∗∗−9.04NS66.25∗∗Punjab Local × R1 B2-2527.04∗∗54.49∗∗33.71∗∗−1.22NS90.44∗∗NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 17
Standard (economic) heterosis of mustard hybrids for yield and its attributed traits.
CrossesTOSSDSSLTWDFIFKranti × R1 B2-2369.00∗∗−7.52∗∗41.53∗∗6.47∗∗17.43∗∗Kranti × R1 B2-866.24∗∗0.12NS7.00NS−0.70NS24.40∗∗Kranti × R1 B2-669.56∗∗−0.25NS31.20∗∗3.13∗∗20.92∗∗Kranti × R1 B2-2972.39∗∗0.54NS18.80∗∗5.82∗∗16.27∗∗Kranti × R1 B2-2581.55∗∗0.37NS18.80∗∗6.47∗∗12.78∗∗JD 6 × R1 B2-2377.63∗∗−3.32NS5.23NS6.55∗∗12.78∗∗JD 6 × R1 B2-870.56∗∗−5.32∗∗11.79NS2.15∗17.43∗∗JD 6 × R1 B2-680.35∗∗−4.12∗8.26NS1.66NS10.45∗∗JD 6 × R1 B2-2981.51∗∗2.35NS9.54NS2.31∗10.45∗∗JD 6 × R1 B2-2584.17∗∗4.31∗17.37∗5.98∗∗9.29∗∗R1 B2-5 × R1 B2-2361.01∗∗−1.35NS21.12∗∗−8.85∗∗25.57∗∗R1 B2-5 × R1 B2-856.27∗∗0.33NS9.80NS−3.31∗∗26.73∗∗R1 B2-5 × R1 B2-677.86∗∗−4.96∗10.81NS1.34NS20.92∗∗R1 B2-5 × R1 B2-2973.95∗∗−8.41∗∗13.19∗2.15∗19.75∗∗R1 B2-5 × R1 B2-2580.95∗∗−8.15∗∗12.41NS6.72∗∗11.61∗∗Varuna × R1 B2-2382.13∗∗−13.85∗∗12.07NS2.40∗11.61∗∗Varuna × R1 B2-877.60∗∗−7.15∗∗0.65NS0.44NS22.08∗∗Varuna × R1 B2-677.35∗∗−6.37∗∗−5.28NS0.85NS15.10∗∗Varuna × R1 B2-2975.32∗∗0.59NS−3.84NS2.07NS10.45∗∗Varuna × R1 B2-2582.29∗∗−6.32∗∗15.19∗6.80∗∗12.78∗∗R1 B2-26 × R1 B2-2386.22∗∗0.23NS30.22∗∗5.00∗∗9.29∗∗R1 B2-26 × R1 B2-872.89∗∗−10.50∗∗24.27∗∗4.43∗∗22.08∗∗R1 B2-26 × R1 B2-684.52∗∗2.68NS23.68∗∗5.49∗∗11.61∗∗R1 B2-26 × R1 B2-2990.90∗∗−7.00∗∗32.10∗∗5.58∗∗10.45∗∗R1 B2-26 × R1 B2-2597.11∗∗−1.72NS19.61∗∗7.04∗∗9.29∗∗T 6342 × R1 B2-2371.41∗∗−2.23NS22.76∗∗5.17∗∗20.92∗∗T 6342 × R1 B2-847.44∗∗−8.67∗∗1.55NS4.60∗∗26.73∗∗T 6342 × R1 B2-671.38∗∗−2.65NS13.60∗5.33∗∗19.75∗∗T 6342 × R1 B2-2984.60∗∗−4.49∗41.36∗∗5.74∗∗12.78∗∗T 6342 × R1 B2-2584.36∗∗3.63NS18.80∗∗6.96∗∗10.45∗∗Punjab Local × R1 B2-2331.48∗∗−6.06∗∗39.68∗∗2.97∗∗38.36∗∗Punjab Local × R1 B2-8−5.22NS−7.36∗∗35.79∗∗2.56∗55.80∗∗Punjab Local × R1 B2-641.19∗∗−1.19NS28.96∗∗3.86∗∗37.19∗∗Punjab Local × R1 B2-2950.11∗∗−7.10∗∗41.77∗∗4.11∗∗31.38∗∗Punjab Local × R1 B2-2570.79∗∗−5.27∗∗10.18NS6.06∗∗22.08∗∗NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 18
Standard (economic) heterosis of mustard hybrids for yield and its attributed traits.
CrossesDFFDMMDYTTBIYKranti × R1 B2-2311.53∗∗−7.14∗∗−15.45∗∗−5.61NS−8.08NSKranti × R1 B2-817.30∗∗−6.55∗∗−17.02∗∗−13.02∗∗20.10∗∗Kranti × R1 B2-614.41∗∗−6.85∗∗−16.24∗∗−8.96∗13.80∗Kranti × R1 B2-2911.53∗∗−7.14∗∗−15.06∗∗−5.70NS−8.07NSKranti × R1 B2-257.68∗−7.14∗∗−13.88∗∗−22.74∗∗−20.84∗∗JD 6 × R1 B2-237.68∗−7.14∗∗−13.88∗∗3.22NS5.01NSJD 6 × R1 B2-811.53∗∗−7.43∗∗−15.84∗∗−6.23NS1.11NSJD 6 × R1 B2-66.72∗−7.14∗∗−13.09∗∗3.40NS−17.59∗∗JD 6 × R1 B2-292.87NS−6.85∗∗−12.70∗∗5.43NS17.48∗∗JD 6 × R1 B2-251.91NS−7.43∗∗−13.09∗∗−21.59∗∗−8.73NSR1 B2-5 × R1 B2-2315.37∗∗−7.43∗∗−18.60∗∗−16.20∗∗−22.57∗∗R1 B2-5 × R1 B2-816.33∗∗−8.02∗∗−19.78∗∗−18.76∗∗−27.64∗∗R1 B2-5 × R1 B2-611.53∗∗−7.43∗∗−17.02∗∗−10.73∗∗−26.70∗∗R1 B2-5 × R1 B2-2910.57∗∗−7.43∗∗−16.63∗∗−6.23NS−22.92∗∗R1 B2-5 × R1 B2-253.84NS−7.43∗∗−13.88∗∗−22.65∗∗−22.68∗∗Varuna × R1 B2-233.84NS−7.43∗∗−13.88∗∗3.93NS−25.99∗∗Varuna × R1 B2-812.49∗∗−8.32∗∗−18.60∗∗−8.61∗−21.79∗∗Varuna × R1 B2-66.72∗−7.73∗∗−15.45∗∗−5.08NS−27.61∗∗Varuna × R1 B2-292.87NS−6.85∗∗−12.70∗∗0.22NS−17.09∗∗Varuna × R1 B2-254.80NS−7.14∗∗−13.88∗∗−22.47∗∗−14.71∗R1 B2-26 × R1 B2-231.91NS−6.85∗∗−12.30∗∗4.81NS2.69NSR1 B2-26 × R1 B2-818.26∗∗−7.14∗∗−17.02∗∗−7.99∗−23.15∗∗R1 B2-26 × R1 B2-69.60∗∗−7.73∗∗−14.27∗∗3.40NS−0.52NSR1 B2-26 × R1 B2-298.64∗∗−7.14∗∗−13.09∗∗4.28NS−12.91∗R1 B2-26 × R1 B2-257.68∗−7.14∗∗−12.70∗∗5.08NS−5.73NST 6342 × R1 B2-2317.30∗∗−7.73∗∗−17.42∗∗−9.23∗3.61NST 6342 × R1 B2-822.10∗∗−7.73∗∗−19.38∗∗−21.06∗∗12.00∗T 6342 × R1 B2-616.33∗∗−6.85∗∗−15.84∗∗−5.61NS−2.60NST 6342 × R1 B2-2910.57∗∗−5.67∗∗−11.91∗∗2.52NS−13.70∗T 6342 × R1 B2-258.64∗∗−6.26∗∗−11.91∗∗−21.06∗∗−9.12NSPunjab Local × R1 B2-2331.72∗∗−7.73∗∗−23.32∗∗−37.57∗∗−31.12∗∗Punjab Local × R1 B2-843.26∗∗−5.97∗∗−26.86∗∗−55.50∗∗−57.74∗∗Punjab Local × R1 B2-627.87∗∗−5.67∗∗−20.17∗∗−32.45∗∗−34.80∗∗Punjab Local × R1 B2-2923.07∗∗−5.67∗∗−18.20∗∗−24.33∗∗0.28NSPunjab Local × R1 B2-2515.37∗∗−7.14∗∗−17.02∗∗−34.13∗∗−9.26NSNS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 19
Standard (economic) heterosis of mustard hybrids for yield and its attributed traits.
CrossesSYHIAIIOICPHCKranti × R1 B2-23−14.87NS13.70NS−29.12∗∗−7.63NS−8.29∗Kranti × R1 B2-830.03∗∗−21.14∗−22.37∗∗−10.24∗−4.84NSKranti × R1 B2-619.04∗−13.00NS−25.74∗∗−2.67NS−6.57∗Kranti × R1 B2-29−14.59NS13.02NS−27.99∗∗−1.66NS−7.71∗Kranti × R1 B2-25−37.90∗∗45.19∗∗−33.62∗∗1.46NS−10.59∗∗JD 6 × R1 B2-23−0.73NS9.44NS−33.62∗∗−4.85NS−10.59∗∗JD 6 × R1 B2-8−1.21NS2.37NS−30.24∗∗−9.32∗−8.86∗∗JD 6 × R1 B2-6−33.22∗∗39.44∗∗−32.49∗∗−2.33NS−10.01∗∗JD 6 × R1 B2-2915.98NS0.09NS−31.37∗∗0.95NS−9.44∗∗JD 6 × R1 B2-25−21.37∗∗28.11∗∗−33.62∗∗1.97NS−7.83∗R1 B2-5 × R1 B2-23−30.26∗∗19.34∗−22.37∗∗−0.22NS−4.84NSR1 B2-5 × R1 B2-8−36.18∗∗23.56∗−22.37∗∗−5.53NS−4.84NSR1 B2-5 × R1 B2-6−38.66∗∗33.64∗∗−25.74∗∗0.20NS−6.57∗R1 B2-5 × R1 B2-29−35.80∗∗34.42∗∗−29.12∗∗0.62NS−8.29∗R1 B2-5 × R1 B2-25−40.86∗∗49.59∗∗−33.62∗∗3.14NS−9.44∗∗Varuna × R1 B2-23−45.64∗∗56.32∗∗−32.49∗∗−3.00NS−10.70Varuna × R1 B2-8−32.74∗∗28.42∗∗−25.74∗∗−10.83∗∗−6.57∗Varuna × R1 B2-6−43.06∗∗44.76∗∗−29.12∗∗−3.59NS−8.29∗Varuna × R1 B2-29−32.36∗∗38.26∗∗−32.49∗∗−2.83NS−10.13∗∗Varuna × R1 B2-25−29.49∗∗35.87∗∗−35.87∗∗−1.57NS−11.74∗∗R1 B2-26 × R1 B2-23−6.37NS16.88NS−36.99∗∗2.72NS−12.43∗∗R1 B2-26 × R1 B2-8−33.60∗∗27.52∗∗−25.74∗∗−0.98NS−6.57∗R1 B2-26 × R1 B2-6−8.76NS15.70NS−35.87∗∗4.41NS−13.00∗∗R1 B2-26 × R1 B2-29−27.01∗∗33.28∗∗−35.87∗∗8.53∗−11.74∗∗R1 B2-26 × R1 B2-25−18.79∗28.09∗∗−36.99∗∗9.46∗−13.58∗∗T 6342 × R1 B2-234.33NS−4.25NS−25.74∗∗−5.87NS−6.57∗T 6342 × R1 B2-822.39∗∗−23.35∗−18.99∗∗−11.76∗∗−3.12NST 6342 × R1 B2-6−6.85NS6.95NS−31.37∗∗−3.34NS−9.44∗∗T 6342 × R1 B2-29−27.29∗∗32.30∗∗−32.49∗∗−2.16NS−8.98∗∗T 6342 × R1 B2-25−22.33∗∗29.61∗∗−38.12∗∗−1.15NS−12.89∗∗Punjab Local × R1 B2-23−30.45∗∗−4.89NS−14.49∗∗−4.52NS−2.08NSPunjab Local × R1 B2-8−59.01∗∗4.12NS−5.49NS−12.52∗∗0.10NSPunjab Local × R1 B2-6−38.66∗∗10.57NS−12.24∗−4.34NS−0.93NSPunjab Local × R1 B2-297.48NS−18.60∗−23.49∗∗−3.26NS−6.68∗Punjab Local × R1 B2-25−15.26NS11.97NS−25.74∗∗−1.99NS−6.57∗NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 20
Standard (economic) heterosis of mustard hybrids for yield and its attributed traits.
CrossesSUCPRCYPPOYPKranti × R1 B2-23−1.34NS−2.19NS0.85NS−6.90NSKranti × R1 B2-80.20NS−1.84NS−8.60∗−18.02∗∗Kranti × R1 B2-61.31NS−2.07NS−4.46NS−7.06NSKranti × R1 B2-29−1.45NS−2.16NS0.25NS−1.45NSKranti × R1 B2-25−3.51NS−2.64NS10.92∗∗12.47∗JD 6 × R1 B2-23−3.51NS−2.66NS10.88∗∗5.44NSJD 6 × R1 B2-8−1.38NS−2.14NS−0.14NS−9.49∗JD 6 × R1 B2-6−3.65NS−2.66NS10.90∗∗8.26NSJD 6 × R1 B2-29−4.07NS−2.72NS13.46∗∗14.48∗∗JD 6 × R1 B2-25−4.14NS−2.83NS12.78∗∗14.96∗∗R1 B2-5 × R1 B2-230.09NS−1.82NS−10.84∗∗−11.08∗R1 B2-5 × R1 B2-80.93NS−1.69NS−13.75∗∗−18.57∗∗R1 B2-5 × R1 B2-6−0.29NS−1.91NS−5.53NS−5.39NSR1 B2-5 × R1 B2-29−1.24NS−2.16NS−0.03NS0.53NSR1 B2-5 × R1 B2-25−3.37NS−2.66NS11.57∗∗15.02∗∗Varuna × R1 B2-23−3.47NS−2.67NS11.61∗∗8.22NSVaruna × R1 B2-8−0.33NS−1.91NS−3.09NS−13.63∗∗Varuna × R1 B2-6−1.31NS−2.14NS1.09NS−2.59NSVaruna × R1 B2-29−3.19NS−2.61NS10.57∗∗7.38NSVaruna × R1 B2-25−17.52∗∗−2.71NS11.78∗∗9.97∗R1 B2-26 × R1 B2-23−3.86NS−2.79NS15.82∗∗18.92∗∗R1 B2-26 × R1 B2-8−0.92NS−1.89NS−5.45NS−6.41NSR1 B2-26 × R1 B2-6−3.40NS−2.69NS11.04∗∗15.88∗∗R1 B2-26 × R1 B2-29−3.65NS−2.66NS12.00∗∗21.49∗∗R1 B2-26 × R1 B2-25−4.17NS−2.89NS16.49∗∗27.45∗∗T 6342 × R1 B2-23−0.78NS−1.89NS−4.32NS−9.97∗T 6342 × R1 B2-81.73NS−1.14NS−17.18∗∗−26.96∗∗T 6342 × R1 B2-6−1.34NS−2.11NS0.52NS−2.88NST 6342 × R1 B2-29−3.05NS−2.52NS10.16∗∗7.72NST 6342 × R1 B2-25−3.51NS−2.76NS13.64∗∗12.29∗Punjab Local × R1 B2-234.84NS0.28NS−36.80∗∗−39.69∗∗Punjab Local × R1 B2-88.61∗1.90NS−57.61∗∗−62.93∗∗Punjab Local × R1 B2-63.83NS−0.18NS−30.53∗∗−33.58∗∗Punjab Local × R1 B2-292.47NS−1.05NS−21.36∗∗−23.95∗∗Punjab Local × R1 B2-25−1.20NS−2.07NS−1.98NS−3.97NSNS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Aside from oil yield heterosis, all 10 crosses showed significant plant height heterosis. R1 B2-26 × R1 B2-25 exhibited the highest oil yield per plant heterosis and a positive plant height heterosis. The total number of primary branches, secondary branches, siliquae on branches, and total siliquae per plant and siliqua length are all factors to consider. The day to first flowering and day to 50% flowering of a plant are used to calculate its seed yield. Days to maturity, stick production per plant, aphid, infestation index, phenol, sugar, and protein contents are all negative heterosis in this cross (Tables16–20). For most of the major characters, including seed yield, Ghosh et al. [53] found that both additive and nonadditive gene actions were critical. Plant height and yield are affected by nonadditive gene action, according to Premlatha et al. [54]. On seed weight and plant height, Gnanasekaran et al. [55] discovered a nonadditive gene effect, whereas Sharma [56] discovered an additive gene effect. Verma et al. [57] found σ2sca variation over σ2gca in barley. Both Borghi et al. [58] and Borghi and Perenzin [59] discovered that 2gca was more essential than σ2sca for most characters. Nonadditive σ2sca is most common in space planting, says Lucken [60]. The disparity in results between research projects may be due to differences in breeding material and genotype conditions.


## 3.1. Genetic Variability among Parents and Hybrids

In comparison with the parents, hybrids were of short duration and taller; produced more primary and secondary branches per plant; had a greater number of siliquae on the main raceme, siliquae on branches, and siliquae per plant; and had a greater weight, oil content, and seed yield per plant. Among the parents, JD6 produces the most oil, while R1 B2-5 has the shortest duration (Table2). In relation to testers, the lines were shorter, with more test weight, and contained a greater quantity of siliquae per plant. The hybrids’ mean values for siliqua length, first flowering day, days to 50% flowering, the yield of the top 10 cm of the main raceme, biological yield per crop, stick yield per crop, aphid infestation content, and phenol content were all within the limits of the parental means. T 6342 × R1 B2-23, R1 B2-26 × R1 B2-29, and Varuna × R1 B2-23 cross-combinations all yielded more oil than the parents on average. Seed number per siliqua, siliqua length, days to first flowering, and phenol content were all same in parents and hybrids. Although genetic diversity and parents’ and hybrids’ mean performance were important parameters for genotypic evaluation, parents with high mean values may not pass this trait onto their hybrids. Using general combining ability (GCA) and specific combining ability (SCA), the effects of these parental and hybrid qualities were calculated.Table 2
Mean performance of lines, testers, and hybrids for yield and its attributed traits in mustard.
GenotypesPHPRBSEBSMRSBTOSSDSSLTWDFIFDFFDMMDYTTBIYSYHIAIIOICPHCSUCPRCYPPOYPLinesKranti154.48.48.147.7131.8179.512.85.04.231.037.2108.077.03.448.834.70.293.437.75.79.720.114.15.3JD 6160.65.75.551.1187.4238.512.73.74.128.734.7113.484.83.850.134.90.313.039.65.89.519.916.36.4R1 B2-5160.65.95.646.9106.5153.313.54.23.831.636.6105.273.63.255.542.70.232.039.05.69.820.112.95.0Varuna131.26.76.548.7128.8177.513.04.44.030.734.7108.377.73.452.238.70.263.139.95.79.720.113.55.4R1 B2-26170.86.66.353.7177.8231.414.54.54.328.732.7109.680.93.848.431.80.351.938.55.99.620.116.66.4T 6342172.56.76.449.7191.9241.613.14.54.330.734.7106.375.63.447.933.60.322.036.25.89.920.314.35.2Punjab Local205.77.97.849.1220.2269.311.44.44.130.834.6123.592.73.026.815.60.452.936.95.49.018.011.24.2Mean (Lines)165.16.86.649.5163.5213.013.04.44.130.335.0110.680.33.447.133.10.322.638.35.79.619.814.15.4TestersR1 B2-23172.57.26.940.5162.6203.113.15.54.235.640.9104.669.02.226.418.20.312.039.85.510.420.88.23.2R1 B2-8170.56.56.838.6105.0143.612.84.63.837.041.0114.777.82.026.918.90.302.034.75.19.819.08.12.8R1 B2-6163.34.44.432.7144.9177.613.94.03.943.548.0110.867.41.322.616.40.281.934.94.910.219.86.22.2R1 B2-29213.55.85.735.1214.9250.012.13.03.940.545.0113.873.21.527.521.10.231.937.04.910.019.76.42.4R1 B2-25192.36.46.440.2203.0243.211.24.03.936.340.8114.478.12.226.517.40.341.937.15.110.019.49.13.4Mean (Testers)182.46.16.037.4166.1203.512.64.24.038.643.1111.773.11.826.018.40.291.936.75.110.119.87.62.8HybridsKranti × R1 B2-23184.710.07.650.5352.6403.111.85.24.433.738.7105.371.73.646.129.70.362.136.65.39.419.516.46.0Kranti × R1 B2-8190.09.57.247.8348.7396.512.84.04.135.740.7106.070.33.360.245.40.252.335.55.59.619.614.95.3Kranti × R1 B2-6200.09.510.249.5354.9404.412.74.94.234.739.7105.771.03.457.141.50.272.238.55.49.719.515.56.0Kranti × R1 B2-29188.59.57.350.2361.0411.212.84.44.333.338.7105.372.03.646.129.80.352.138.95.49.419.516.36.3Kranti × R1 B2-25197.79.58.753.2379.9433.012.84.44.432.337.3105.373.02.939.721.70.452.040.25.29.219.418.07.2JD 6 × R1 B2-23190.06.36.737.1386.6423.712.33.94.432.337.3105.373.03.952.734.60.342.037.75.29.219.418.06.8JD 6 × R1 B2-8185.36.43.750.7356.1406.812.14.14.233.738.7105.071.33.550.734.50.322.135.95.39.419.516.25.8JD 6 × R1 B2-6185.09.39.253.5376.7430.212.24.04.231.737.0105.373.73.941.323.30.442.038.75.29.219.418.07.0JD 6 × R1 B2-29174.99.69.653.7379.2432.913.04.14.231.735.7105.774.04.058.940.50.312.040.05.39.219.418.47.4JD 6 × R1 B2-25194.09.610.153.6385.6439.313.34.44.331.335.3105.073.73.045.827.40.402.040.45.39.119.418.37.4R1 B2-5 × R1 B2-23181.27.66.647.5336.5384.012.64.53.736.040.0105.069.03.238.824.30.372.339.55.59.619.614.55.7R1 B2-5 × R1 B2-8176.26.96.446.5326.2372.712.84.14.036.340.3104.368.03.136.322.30.392.337.45.59.619.614.05.2R1 B2-5 × R1 B2-6170.07.56.649.0375.2424.212.14.14.134.738.7105.070.33.436.821.40.422.239.75.49.519.615.46.1R1 B2-5 × R1 B2-29177.58.97.050.5364.4414.911.74.24.234.338.3105.070.73.538.722.40.422.139.85.39.419.516.36.5R1 B2-5 × R1 B2-25181.59.36.653.5378.1431.611.74.24.432.036.0105.073.02.938.820.60.472.040.85.39.219.418.17.4Varuna × R1 B2-23159.97.87.553.5380.9434.411.04.24.232.036.0105.073.03.937.119.00.492.038.45.29.219.418.17.0Varuna × R1 B2-8186.36.87.449.0374.6423.611.83.74.135.039.0104.069.03.539.223.50.402.235.35.49.519.615.85.6Varuna × R1 B2-6201.07.27.550.5372.5423.011.93.54.133.037.0104.771.73.636.319.90.452.138.25.39.419.516.46.3Varuna × R1 B2-29195.78.37.453.0365.2418.212.83.64.231.735.7105.774.03.841.623.60.432.038.55.29.219.418.06.9Varuna × R1 B2-25204.09.47.453.3381.5434.811.94.34.432.336.3105.373.02.942.824.60.431.939.05.17.919.418.27.1R1 B2-26 × R1 B2-23203.28.310.454.5389.7444.212.84.84.331.335.3105.774.34.051.532.70.371.940.75.19.219.418.87.7R1 B2-26 × R1 B2-8200.34.95.549.5362.9412.411.44.64.335.041.0105.370.33.538.523.20.402.239.25.49.519.615.46.0R1 B2-26 × R1 B2-6201.68.69.453.5386.6440.113.14.64.332.038.0104.772.73.949.931.80.361.941.35.09.219.418.17.5R1 B2-26 × R1 B2-29192.78.88.353.5401.8455.311.84.94.331.737.7105.373.73.943.725.50.421.943.05.19.219.418.27.8R1 B2-26 × R1 B2-25192.99.711.454.8415.3470.112.54.44.431.337.3105.374.04.047.328.30.401.943.35.09.119.418.98.2T 6342 × R1 B2-23203.36.79.349.2359.7408.812.54.64.334.740.7104.770.03.452.036.40.302.237.35.49.519.615.65.8T 6342 × R1 B2-8180.26.88.446.5305.2351.711.63.84.336.342.3104.768.33.056.242.70.242.434.95.69.719.713.54.7T 6342 × R1 B2-6199.97.08.451.0357.8408.812.44.24.334.340.3105.771.33.648.832.50.332.038.35.39.419.516.36.3T 6342 × R1 B2-29202.29.58.753.1387.2440.312.25.24.332.338.3107.074.73.943.325.40.412.038.75.39.319.417.96.9T 6342 × R1 B2-25204.39.59.453.5386.2439.713.24.44.431.737.7106.374.73.045.627.10.411.839.15.19.219.418.57.2Punjab Local × R1 B2-23200.07.84.541.5272.1313.612.05.24.239.745.7104.765.02.434.524.30.302.537.85.710.020.010.33.9Punjab Local × R1 B2-8163.65.34.937.0189.1226.111.85.04.244.749.7106.762.01.721.214.30.332.834.65.810.420.36.92.4Punjab Local × R1 B2-6180.36.97.643.5293.3336.812.64.84.239.344.3107.067.72.632.721.40.352.637.95.79.919.911.34.3Punjab Local × R1 B2-29187.17.28.346.5311.5358.011.85.34.337.742.7107.069.32.950.337.50.252.338.35.49.819.712.84.9Punjab Local × R1 B2-25204.08.87.350.5356.9407.412.14.14.335.040.0105.370.32.545.529.60.352.238.85.49.419.515.96.2Mean (Hybrids)189.78.17.849.8357.5407.312.34.44.234.139.1105.471.23.344.228.10.372.138.65.39.419.516.16.2Grand mean185.37.77.448.5308.2356.712.44.44.234.038.9106.872.83.242.727.80.352.238.45.49.519.614.95.8SE (±)5.60.30.43.67.57.30.20.20.00.71.10.81.10.12.82.80.030.21.60.20.40.30.60.3CD (P=0.0511.10.60.87.115.014.60.50.50.11.32.11.62.10.35.65.60.060.33.10.40.70.71.20.6PH: plant height (cm), PRB: number of total primary branches per plant, SEB: number of total secondary branches per plant, SMR: number of siliquae on the main raceme, SB: number of siliquae on branches, TOS: total number of siliquae per plant, SDS: no. seeds per siliqua, SL: length of siliqua (cm), TW: test weight (g), DFIF: days to 1st flowering, DFF: days to 50% flowering, DM: days to maturity∗∗, MD: duration of maturity (i.e., duration from 1st flowering to maturity), YTT: yield of the top 10 cm of the main raceme (g), BIY: biological yield per plant (g), SY: stick yield per plant(g), HI: harvest index, AII: aphid infestation index, OIC: oil content (%), PHC: phenol content (mg/g), SUC: sugar content (mg/g), PRC: protein content (%), YPP: yield per plant (g), OYP: oil yield per plant (g).The analysis of variance disclosed that all features under examination had a significant genotype effect. This demonstrates the existence of adequate genetic diversity among lines, testers, and hybrids and enables a more thorough evaluation of the general combining ability studies (Tables3–5). Combining ability analysis is commonly used in self-pollinated crops to identify suitable parents and, more specifically, a cross with a higher possibility for producing desirable recombinant lines. The combining ability study demonstrated that the mean squares for all twenty-four yield and quality features were highly significant among parents, indicating the existence of genetic diversity between the parents (Table 2). The hybrids differed significantly for all parameters, demonstrating genetic diversity, except for days to maturity and protein content, which have modest variability. For all parameters except siliqua length and stick yield per plant, the variation due to parents versus hybrids was remarkable, showing the presence of significant variations between the crosses [36].Table 3
Analysis of variance for combining ability effects of yield and its attributed traits for parents and hybrids in mustard.
Source of variationdfPHPRBSEBSMRSBTOSSDSSLReplication2109.371NS0.334NS0.242NS6.133NS59.114NS56.604NS0.089NS0.091NSGenotypes46858.6776.553∗∗8.257∗∗94.504∗∗26824.021∗∗28792.752∗∗1.499∗∗0.794∗∗Parents111522.963∗∗3.258∗∗3.026∗∗138.196∗∗4967.440∗∗5230.693∗∗2.682∗∗1.220∗∗Parents vs crosses18065.407∗∗70.451∗∗54.977∗∗769.322∗∗997557.780∗∗1053732.616∗∗8.754∗∗0.094NSCrosses34431.798∗∗5.740∗∗8.576∗∗60.520∗∗5344.275∗∗6270.481∗∗0.903∗∗0.677∗∗Lines6805.998NS7.777∗14.400∗131.694∗∗17861.705∗∗20937.150∗∗0.995NS1.900∗∗Testers4524.105NS23.418∗∗20.480∗150.066∗∗10247.879∗∗12689.890∗∗0.827NS0.666NSLine × testers24322.864∗∗2.284∗∗5.136∗∗27.802NS1397.650∗∗1533.912∗∗0.892∗∗0.373∗∗Error9246.6310.1160.22119.11885.42480.6910.0910.091NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 4
Analysis of variance for combining ability effects of yield and its attributed traits for parents and hybrids in mustard.
Source of variationdfTWDFIFDFFDMMDYTTBIYSYReplication20.002NS0.178NS1.012NS1.114NS2.151NS0.004NS2.297NS3.827NSGenotypes460.084∗∗34.951∗∗37.461∗∗40.109∗∗71.908∗∗1.401289.527∗∗194.535∗∗Parents110.107∗∗68.677∗∗67.452∗∗84.613∗∗138.357∗∗2.373∗∗502.125∗∗295.775∗∗Parents vs crosses10.873∗∗4.177∗12.014∗∗860.857∗∗984.963∗∗8.398∗∗916.621∗∗30.804NSCrosses340.054∗∗24.945∗∗28.506∗∗1.571NS23.556∗∗0.881∗∗202.301∗∗166.596∗∗Lines60.105∗90.022∗∗104.200∗∗2.905NS70.327∗∗3.304∗∗524.668∗365.777∗Testers40.127∗∗55.938∗∗59.65∗∗71.929NS68.938∗∗1.593∗∗31.093NS52.209NSLinex testers240.029∗∗3.510∗∗4.390∗∗1.179NS4.299∗∗0.156∗∗150.244∗∗135.866∗∗Error920.0030.6911.7201.0091.7320.03411.89912.008NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 5
Analysis of variance for combining ability effects of yield and its attributed traits for parents and hybrids in mustard.
Source of variationdfHIAIIOICPHCSUCPRCYPPOYPReplication20.001NS0.037NS1.273NS0.010NS0.444NS0.014NS0.340NS0.019NSGenotypes460.014∗∗0.360∗∗11.824∗∗0.187∗∗0.511∗∗0.485∗∗38.431∗∗7.065∗∗Parents110.011∗∗0.959∗∗9.648∗∗0.404∗∗0.370NS1.529∗∗41.377∗∗6.748∗∗Parents vs crosses10.112∗∗1.132∗∗27.862∗∗0.313∗4.681∗∗1.540∗∗590.205∗∗99.128∗∗Crosses340.012∗∗0.143∗∗12.057∗∗0.114∗∗0.434∗∗0.116NS21.249∗∗4.460∗∗Lines60.031∗∗0.457∗∗30.566∗∗0.340∗∗1.084∗∗0.404∗∗73.904∗∗14.012∗∗Testers40.019∗0.38∗∗052.368∗∗0.281∗∗1.120∗∗0.224∗∗50.064∗∗14.486∗∗Linex testers240.007∗∗0.026NS0.711NS0.029NS0.157NS0.027NS3.283∗∗0.400∗∗Error920.0010.0393.6620.0540.2110.1710.5050.141NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Except for plant height, seed number per siliqua, and days to maturity, the lines showed significant differences in all studied characteristics, whereas the testers showed significant differences in the majority of the studied characteristics, except for plant height, seed number per siliqua, siliqua length, day to maturity, and biological and stick yield per plant. Except for plant height, seed number per siliqua, days to maturity, siliqua length, biological and stick yield per crop, additive variance predominated for the majority of the parameters [37].Due to the average squares, the line × tester interaction constituents were also remarkable for all yield components, specifying that merging ability played a significant role in the declaration of these traits and served as a direct test for the significance of dominant or nonadditive variance for the characters under study. However, the same was not found for certain less significant characters such as the total number of siliquae on the main raceme, days to maturity, aphid infestation index, oil content, and other biochemical components such as phenol, sugar, and protein.


## 3.2. General and Specific Combining Abilities

R1 B2-26 was the best combiner for oil yield/plant among the seven lines, although Varuna and JD6 both had significant GCA values. Punjab Local, on the other hand, exhibited a significantly negative GCA, while Kranti, R1 B2-5, and T 6342 had nonsignificant GCAs (Tables6 and 7). This means that by employing R1 B2-26, Varuna, and JD6 as parents, oil yield/plant may be increased; for instance, the number of primary and secondary branches on a plant, the number of siliquae on a raceme, the number of seeds on a raceme, the number of seeds per siliquae on a plant, the number of seeds per siliquae on a plant, the number of seeds per siliquae on a plant, the number of seeds per siliquae on a plant, and the number of seeds per siliquae on a plant. Additionally, R1 B2-26 demonstrated statistically significant GCA values for plant height, secondary branch count, siliqua count (main raceme, branches, and total), siliqua length, weight, maturity duration, biological yield (top 10 cm of the main raceme), harvest index (oil content), and seed production per plant.Table 6
General combining ability (GCA) effects of parents on yield and its attributed traits in mustard.
ParentsPHPRBSEBSMRSBTOSSDSSLTWDFIFDFFDMLines (gi)Kranti2.48NS1.46∗∗0.40∗∗0.40NS1.94NS2.34NS0.29∗∗0.19∗0.03NS−0.20NS−0.07NS0.15NSJD 6−3.84∗0.10NS0.09NS−0.11NS19.36∗∗19.26∗∗0.30∗∗−0.29∗∗0.01NS−2.00∗∗−2.27∗∗−0.11NSR1 B2-5−12.41∗∗−0.09NS−1.14∗∗−0.44NS−1.37NS−1.81NS−0.11NS−0.18∗−0.16∗∗0.53∗−0.40NS−0.51NSVaruna−0.32NS−0.23∗∗−0.36∗∗2.03NS17.45∗∗19.49∗∗−0.38∗∗−0.54−0.04∗∗−1.33∗∗−2.27∗∗−0.45NSR1 B2-268.46∗∗−0.08NS1.23∗∗3.33∗∗33.78∗∗37.11∗∗0.04NS0.29∗∗0.08∗∗−1.87∗∗−1.20∗∗−0.11NST 63428.29∗∗−0.23∗∗1.050.82NS1.73NS2.55NS0.09NS0.05NS0.08∗∗−0.27NS0.80∗0.29NSPunjab Local−2.68NS−0.92∗∗−1.25∗∗−6.04∗∗−72.91∗∗−78.94∗∗−0.23∗∗0.48∗∗0.01NS5.13∗∗5.40∗∗0.75∗∗SE (gi)1.760.090.121.132.392.320.080.080.010.210.340.26Tester (gj)R1 B2-23−0.80NS−0.35∗∗−0.28∗∗−2.15∗−3.47NS−5.62∗∗−0.16∗0.24∗∗−0.03∗∗0.10NS0.03NS−0.29NSR1 B2-8−6.56∗∗−1.49∗∗−1.57∗∗−3.12∗∗−34.22∗∗−37.35∗∗−0.24∗∗−0.20∗∗−0.09∗∗2.53∗∗2.60∗∗−0.24NSR1 B2-61.42NS−0.14NS0.63∗∗0.24NS2.09NS2.33NS0.15∗−0.09NS−0.02NS0.10NS0.22NS0.05NSR1 B2-29−1.31NS0.71∗∗0.29∗∗1.66NS9.72∗∗11.38∗∗0.03NS0.13∗0.02NS−0.90∗∗−0.92∗∗0.48∗R1 B2-257.24∗∗1.26∗∗0.93∗∗3.37∗∗25.88∗∗29.25∗∗0.22∗∗−0.08NS0.12∗∗−1.85∗∗−1.92∗∗0.00NSSE (gj)1.490.070.100.952.021.960.070.070.010.180.290.22NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 7
General combining ability (GCA) effects of parents on yield and its attributed traits in mustard.
ParentsMDYTTBIYSYHIAIIOICPHCSUCPRCYPPOYPLines (gi)Kranti0.35NS0.02NS5.67∗∗5.54∗∗−0.03∗∗0.01NS−0.68NS0.02NS0.06NS−0.03NS0.13NS−0.08NSJD 61.89∗∗0.32∗∗5.71∗∗3.99∗∗−0.01NS−0.12∗−0.12NS−0.08NS−0.17NS−0.12NS1.72∗∗0.62∗∗R1 B2-5−1.05∗∗−0.12∗−6.30∗∗−5.86∗∗0.04∗∗0.05NS0.82NS0.07NS0.08NS−0.01NS−0.44∗−0.06NSVaruna0.89∗0.20∗∗−4.77∗∗−5.97∗∗0.07∗∗−0.09NS−0.77NS−0.09NS−0.34∗∗−0.08NS1.20∗∗0.31∗∗R1 B2-261.75∗∗0.51∗∗2.01∗0.22NS0.02∗−0.18∗∗2.87∗∗−0.20∗∗−0.15NS−0.11NS1.79∗∗1.19∗∗T 63420.550.03NS5.00∗∗4.74∗∗−0.03∗∗−0.03NS−0.96NS−0.01NS0.02NS−0.01NS0.26NS−0.06NSPunjab Local−4.38∗∗−0.95∗∗−7.32∗∗−2.66∗∗−0.06∗∗0.35∗∗−1.15∗0.28∗∗0.51∗∗0.36−4.66∗∗−1.92∗∗SE (gi)0.340.050.890.890.010.050.490.060.120.110.180.10Tester (gj)R1 B2-23−0.39NS0.13∗∗0.51NS0.64NS−0.01NS0.01NS−0.36NS0.01NS0.04NS0.01NS−0.13NS−0.13NSR1 B2-8−2.77∗∗−0.27∗∗−0.97NS1.32NS−0.04∗∗0.20∗∗−2.50∗∗0.18∗∗0.27∗∗0.15NS−2.29∗∗−1.24∗∗R1 B2-6−0.06NS0.14∗∗−0.90NS−0.67NS0.00NS0.02NS0.30NS0.01NS0.09NS0.01NS−0.23NS−0.06NSR1 B2-291.37∗∗0.31∗∗1.90∗1.16NS0.00NS−0.06NS0.97∗−0.06NS−0.04NS−0.05NS0.75∗∗0.43∗∗R1 B2-251.85∗∗−0.31∗∗−0.54NS−2.45∗∗0.04∗∗−0.17∗∗1.60∗∗−0.14∗∗−0.36∗∗−0.13NS1.91∗∗1.00∗∗SE (gi)0.290.040.750.760.010.040.420.050.100.090.160.08NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.For JD6 and Varuna, GCA values for maturity, the top 10 cm of the main raceme yield, and yield per plant were all considerably positive. However, both had a negative impact of GCA on siliqua length, first flowering day, and 50% flowering days. By reducing the height of the plant and aphid infection index, JD6 caused dwarfism and aphid resistance. For days to maturity, both JD6 and Varuna showed high GCA values, indicating an unfavorable increase in crop length. Only Punjab Local had the required significantly negative GCA value for oil yield per plant and plant height. Utilizing this line to enhance the character should be done with caution since it has the potential to significantly reduce the yield.R1 B2-25 and R1 B2-29 had significant positive GCA values for oil yield/plant, whereas R1 B2-8 had significant negative GCA values. When R1 B2-25 and R1 B2-29 are used as parents, the character is enhanced; however, when R1 B2-8 is used as a parent, the oil yield/plant is decreased. Other yield-related characteristics such as plant height, number of primary and secondary branches per plant, number of siliquae on the main raceme, number of siliquae on branches, number of seeds per siliqua, weight, maturity duration, harvest index, oil content, and seed yield per plant all showed significant positive GCA values. R1 B2-29 had a significant positive correlation between the total number of primary and secondary branches on a plant, siliquae on branches, total siliqua per plant, length of siliqua, days to maturity, maturity duration, yield of the top 10 cm of the main raceme, biological yield per plant, oil content, and seed yield per plant. R1 B2-8 has a substantial negative GCA when compared with plant height. The number of major and secondary branches, the number of siliquae on the main raceme and on branches, the number of siliquae per plant, the number of seeds per siliqua, the length of siliqua (cm), and the weight of siliqua are all factors to consider. Despite the presence of dwarfing genes in R1 B2-8, utilizing it to improve features should be done with caution since adopting this line may eventually result in a considerable drop in yield and other yield-related qualities like aphid resistance.However, in JD6, R1 B2-26 between the lines can be chosen as an expected parent for increasing oil yield/plant, secondary branch number per plant, siliqua on the main raceme, siliqua on branches, total siliqua per plant, siliqua length, weight, top 10 cm of the main raceme yield, biological yield per crop, and harvest.R1 B2-25 may be considered as a possible parent to enhance oil yield, aphid resistance, and seed size among the testers, as evidenced by a significant positive GCA for test weight, although it may delay maturity. This parent, on the other hand, may boost the amount of oil, phenol, and sugar. Despite having a strong GCA influence on oil content, R1 B2-29 is a poor general combiner.While SCA effects do not directly benefit self-pollinated crops, ideal hybrids should provide transgressive segregants that can be selected as potent homozygous lines.Punjab Local × R1 B2-25 and Varuna × R1 B2-23 were the only crossings with significantly positive SCA values for oil yield/plant (Tables8–12). On the other hand, three crossings (Punjab Local × R1 B2-8, Varuna × R1 B2-25, and JD6 × R1 B2-25) had considerably negative SCA for plant yield. Plant height, number of siliquae on branches, number of siliquae per plant, maturity, yield of the top 10 cm of the main raceme, biological yield per plant, stick yield per plant, and seed yield per plant have all exhibited a significant positive SCA effect in the cross Punjab Local × R1 B2-8. This hybrid, on the other hand, showed significant SCA impacts on plant height and seed yield per siliqua.Table 8
Specific combining ability (SCA) effects of mustard hybrids on yield and its attributed traits.
HybridsPHPRBSEBSMRSBKranti × R1 B2-23−6.704NS0.717∗∗−0.344NS2.419NS−3.344NSKranti × R1 B2-84.391NS1.389∗∗0.587∗0.721NS23.478∗∗Kranti × R1 B2-66.410NS0.070NS1.382∗∗−0.969NS−6.575NSKranti × R1 B2-29−2.390NS−0.778∗∗−1.208∗∗−1.731NS−8.132NSKranti × R1 B2-25−1.709NS−1.397∗∗−0.418NS−0.440NS−5.427NSJD 6 × R1 B2-234.950NS−1.556∗∗−0.897∗∗−10.461∗∗13.183∗JD 6 × R1 B2-86.045NS−0.318NS−2.600∗∗4.058NS13.521∗JD 6 × R1 B2-6−2.303NS1.163∗∗0.696∗3.534NS−2.265NSJD 6 × R1 B2-29−9.603∗0.615∗∗1.472∗∗2.339NS−7.355NSJD 6 × R1 B2-250.911NS0.096NS1.329∗∗0.530NS−17.084∗∗R1 B2-5 × R1 B2-234.756NS−0.103NS0.243NS0.252NS−16.097∗∗R1 B2-5 × R1 B2-85.451NS0.335NS1.324∗∗0.221NS4.358NSR1 B2-5 × R1 B2-6−8.696∗−0.384−0.681∗−0.636NS17.039∗∗R1 B2-5 × R1 B2-291.504NS0.169NS0.062NS−0.564NS−1.419NSR1 B2-5 × R1 B2-25−3.015NS−0.017NS−0.9480.726NS−3.880NSVaruna × R1 B2-23−28.704∗∗0.210NS0.316NS3.786NS9.443NSVaruna × R1 B2-83.525NS0.415∗1.514∗∗0.255NS33.898∗∗Varuna × R1 B2-610.177∗−0.604∗∗−0.524NS−1.602NS−4.521NSVaruna × R1 B2-297.610NS−0.285NS−0.315NS−0.531NS−19.479∗∗Varuna × R1 B2-257.391NS0.263NS−0.991∗∗−1.907NS−19.340∗∗R1 B2-26 × R1 B2-235.823NS0.6241.663∗∗3.486NS1.883NSR1 B2-26 × R1 B2-88.685∗−1.705−1.906∗∗−0.545NS5.838NSR1 B2-26 × R1 B2-62.037NS0.643−0.244NS0.098NS−6.748NSR1 B2-26 × R1 B2-29−4.096NS0.062NS−1.001∗∗−1.331NS0.861NSR1 B2-26 × R1 B2-25−12.449∗∗0.376NS1.489∗∗−1.707NS−1.834NST 6342 × R1 B2-236.090NS−0.856∗∗0.743∗∗0.666NS3.930NST 6342 × R1 B2-8−11.249∗∗0.349NS1.107∗∗−1.032NS−19.816∗∗T 6342 × R1 B2-60.504NS−0.737∗∗−1.064∗∗0.111NS−3.535NST 6342 × R1 B2-295.537NS0.915∗∗−0.455NS0.782NS18.275∗∗T 6342 × R1 B2-25−0.882NS0.330NS−0.331NS−0.527NS1.146NSPunjab Local × R1 B2-2313.790∗∗0.964∗∗−1.724∗∗−0.148NS−8.997NSPunjab Local × R1 B2-8−16.849∗∗−0.465∗−0.026NS−3.679NS−61.276∗∗Punjab Local × R1 B2-6−8.130∗−0.150NS0.436NS−0.536NS6.605NSPunjab Local × R1 B2-291.437NS−0.698∗∗1.445∗∗1.036NS17.248∗∗Punjab Local × R1 B2-259.751∗0.350NS−0.131NS3.326NS46.420∗∗SE (±)3.9430.1970.2722.5245.336NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 9
Specific combining ability (SCA) effects of mustard hybrids on yield and its attributed traits.
HybridsTOSSDSSLTWDFIFKranti × R1 B2-23−0.925NS−0.625∗∗0.434∗0.123∗∗−0.371NSKranti × R1 B2-824.199∗∗0.429∗−0.414∗−0.115∗∗−0.800NSKranti × R1 B2-6−7.544NS−0.012NS0.376∗−0.025NS0.629NSKranti × R1 B2-29−9.863NS0.209NS−0.305NS0.049NS0.295NSKranti × R1 B2-25−5.868NS−0.001NS−0.090NS−0.031NS0.248NSJD 6 × R1 B2-232.722NS−0.106NS−0.429∗0.147∗∗0.095NSJD 6 × R1 B2-817.579∗∗−0.280NS0.247NS0.022NS−1.000∗JD 6 × R1 B2-61.270NS−0.522∗∗0.008NS−0.065∗−0.571NSJD 6 × R1 B2-29−5.016NS0.423∗−0.165NS−0.074∗0.429NSJD 6 × R1 B2-25−16.554∗∗0.484∗∗0.340NS−0.030NS1.048∗R1 B2-5 × R1 B2-23−15.845∗∗0.564∗∗0.047NS−0.315∗∗1.229∗R1 B2-5 × R1 B2-84.579NS0.859∗∗0.061NS−0.032NS−0.867NSR1 B2-5 × R1 B2-616.403∗∗−0.210NS−0.010NS0.091∗∗−0.105NSR1 B2-5 × R1 B2-29−1.983−0.529∗∗−0.1420.088∗∗0.562R1 B2-5 × R1 B2-25−3.154NS−0.684∗∗0.044NS0.168∗∗−0.819NSVaruna × R1 B2-2313.229∗−0.760∗∗0.072NS0.027NS−0.905NSVaruna × R1 B2-834.152∗∗0.175NS0.082NS0.002NS−0.333NSVaruna × R1 B2-6−6.124NS−0.121NS−0.246NS−0.048NS0.095NSVaruna × R1 B2-29−20.010∗∗0.887−0.414∗−0.034NS−0.238NSVaruna × R1 B2-25−21.248∗∗−0.1810.507∗∗0.053NS1.381∗∗R1 B2-26 × R1 B2-235.369NS0.606∗∗−0.079NS0.011NS−1.038∗R1 B2-26 × R1 B2-85.292NS−0.680∗∗0.133NS0.043NS0.200NSR1 B2-26 × R1 B2-6−6.650NS0.605∗∗0.004NS0.019NS−0.371NSR1 B2-26 × R1 B2-29−0.470NS−0.507∗∗0.095NS−0.013NS0.295NSR1 B2-26 × R1 B2-25−3.541NS−0.024NS−0.153NS−0.060NS0.914NST 6342 × R1 B2-234.595NS0.244NS−0.120NS0.015NS0.695NST 6342 × R1 B2-8−20.848∗∗−0.495∗∗−0.473∗∗0.048NS−0.067NST 6342 × R1 B2-6−3.424NS−0.123NS−0.134NS0.011NS0.362NST 6342 × R1 B2-2919.057∗∗−0.235NS0.674−0.008NS−0.638NST 6342 × R1 B2-250.619NS0.609∗∗0.053NS−0.065∗−0.352NSPunjab Local × R1 B2-23−9.145NS0.077NS0.075NS−0.007NS0.295NSPunjab Local × R1 B2-8−64.954∗∗−0.008NS0.364∗0.032NS2.867∗∗Punjab Local × R1 B2-66.070NS0.383∗0.003NS0.018NS−0.038NSPunjab Local × R1 B2-2918.284∗∗−0.249NS0.257NS−0.008NS−0.705NSPunjab Local × R1 B2-2549.746∗∗−0.204NS−0.700∗∗−0.034NS−2.419∗∗SE (±)5.1860.1740.1740.0310.480NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 10
Specific combining ability (SCA) effects of mustard hybrids on yield and its attributed traits.
HybridsDFFDMMDYTTBIYKranti × R1 B2-23−0.362NS0.086NS0.457NS0.078NS−4.247∗Kranti × R1 B2-8−0.933NS0.705NS1.505NS0.200NS11.362∗∗Kranti × R1 B2-60.448NS0.086NS−0.543NS−0.052NS8.124∗∗Kranti × R1 B2-290.590NS−0.676NS−0.971NS−0.103NS−5.642∗∗Kranti × R1 B2-250.257NS−0.200NS−0.448NS−0.122NS−9.597∗∗JD 6 × R1 B2-230.505NS0.352NS0.257NS0.107NS2.279NSJD 6 × R1 B2-8−0.733NS−0.029NS0.971NS0.152NS1.801NSJD 6 × R1 B2-6−0.019NS0.019NS0.590NS0.110NS−7.650∗∗JD 6 × R1 B2-29−0.210NS−0.076NS−0.505NS0.013NS7.134∗∗JD 6 × R1 B2-250.457NS−0.267NS−1.314NS−0.383∗∗−3.564NSR1 B2-5 × R1 B2-231.305NS0.419NS−0.810NS−0.182NS0.463NSR1 B2-5 × R1 B2-8−0.933NS−0.295NS0.571NS0.123NS−0.598NSR1 B2-5 × R1 B2-6−0.219NS0.086NS0.190NS0.021NS−0.206NSR1 B2-5 × R1 B2-290.590NS−0.343NS−0.905NS0.017NS−1.112NSR1 B2-5 × R1 B2-25−0.743NS0.133NS0.952NS0.021NS1.453NSVaruna × R1 B2-23−0.829NS0.352NS1.257NS0.256∗−2.790NSVaruna × R1 B2-8−0.400NS−0.695NS−0.362NS0.185NS0.799NSVaruna × R1 B2-6−0.019NS−0.314NS−0.410NS−0.087NS−2.199NSVaruna × R1 B2-29−0.210NS0.257NS0.495NS−0.061NS0.275NSVaruna × R1 B2-251.457NS0.400NS−0.981NS−0.293∗∗3.916NSR1 B2-26 × R1 B2-23−2.562∗∗0.686NS1.724∗−0.024NS4.816∗R1 B2-26 × R1 B2-80.533NS0.305NS0.105NS−0.106NS−6.661∗∗R1 B2-26 × R1 B2-6−0.086NS−0.648NS−0.276NS−0.081NS4.608∗R1 B2-26 × R1 B2-290.724NS−0.410NS−0.705NS−0.222∗−4.402∗R1 B2-26 × R1 B2-251.390NS0.067−0.848NS0.433∗∗1.640NST 6342 × R1 B2-230.771NS−0.714−1.410NS−0.071NS2.287NST 6342 × R1 B2-8−0.133NS−0.762NS−0.695NS−0.116NS7.976∗∗T 6342 × R1 B2-60.248NS−0.048NS−0.410NS0.062NS0.575NST 6342 × R1 B2-29−0.610NS0.857NS1.495NS0.195NS−7.791∗∗T 6342 × R1 B2-25−0.276NS0.667NS1.019NS−0.071NS−3.046NSPunjab Local × R1 B2-231.171NS−1.181∗−1.476NS−0.163NS−2.807NSPunjab Local × R1 B2-82.600∗∗0.771NS−2.095∗∗−0.438∗∗−14.678∗∗Punjab Local × R1 B2-6−0.352NS0.819NS0.857NS0.027NS−3.252NSPunjab Local × R1 B2-29−0.876NS0.390NS1.095NS0.160NS11.538∗∗Punjab Local × R1 B2-25−2.543∗∗−0.800NS1.619∗0.414∗∗9.200∗∗SE (±)0.7570.5800.7600.1061.992NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 11
Specific combining ability (SCA) effects of mustard hybrids on yield and its attributed traits.
HybridsSYHIAIIOICPHCKranti × R1 B2-23−4.552∗0.029NS−0.052NS−1.016NS−0.045NSKranti × R1 B2-810.433∗∗−0.050∗−0.038NS0.089NS−0.018NSKranti × R1 B2-68.586∗∗−0.068∗∗0.038NS0.288NS0.049NSKranti × R1 B2-29−4.971∗0.016NS0.057NS0.017NS0.050NSKranti × R1 B2-25−9.495∗∗0.074∗∗−0.005NS0.622NS−0.037NSJD 6 × R1 B2-231.934NS−0.010NS−0.052NS−0.483NS−0.077NSJD 6 × R1 B2-81.087NS−0.003NS−0.138NS−0.111NS−0.150NSJD 6 × R1 B2-6−8.094∗∗0.070∗∗−0.029NS−0.145NS−0.050NSJD 6 × R1 B2-297.249∗∗−0.051∗0.090NS0.484NS0.051NSJD 6 × R1 B2-25−2.175NS−0.006NS0.129NS0.255NS0.225NSR1 B2-5 × R1 B2-231.488NS−0.030NS0.114NS0.417NS0.108NSR1 B2-5 × R1 B2-8−1.260NS0.013NS−0.071NS0.455NS−0.064NSR1 B2-5 × R1 B2-6−0.141NS0.001NS0.005NS−0.078NS0.002NSR1 B2-5 × R1 B2-29−0.965NS0.006NS−0.010NS−0.583NS−0.030NSR1 B2-5 × R1 B2-250.878NS0.010NS−0.038NS−0.211NS−0.017NSVaruna × R1 B2-23−3.772NS0.059∗∗−0.052NS0.904NS−0.076NSVaruna × R1 B2-80.047NS0.001NS−0.038NS−0.058NS−0.008NSVaruna × R1 B2-6−1.568NS0.009NS0.038NS0.008NS0.058NSVaruna × R1 B2-290.342NS−0.009NS0.024NS−0.363NS0.019NSVaruna × R1 B2-254.951∗−0.059∗∗0.029NS−0.491NS0.006NSR1 B2-26 × R1 B2-233.734NS−0.013NS−0.092NS−0.469NS−0.061NSR1 B2-26 × R1 B2-8−6.447∗∗0.049∗0.055NS0.202NS0.106NSR1 B2-26 × R1 B2-64.206∗−0.031NS−0.069NS−0.465NS−0.100NSR1 B2-26 × R1 B2-29−3.985∗0.027NS0.017NS0.497NS0.041NSR1 B2-26 × R1 B2-252.491NS−0.032NS0.089NS0.235NS0.014NST 6342 × R1 B2-232.948NS−0.029NS0.094NS−0.036NS0.090NST 6342 × R1 B2-88.567∗∗−0.060∗∗0.109NS−0.231NS0.117NST 6342 × R1 B2-60.352NS−0.008NS−0.082NS0.302NS−0.083NST 6342 × R1 B2-29−8.605∗∗0.074∗∗−0.030NS0.097NS0.011NST 6342 × R1 B2-25−3.262NS0.023NS−0.091NS−0.131NS−0.135NSPunjab Local × R1 B2-23−1.779NS−0.007NS0.041NS0.683NS0.062NSPunjab Local × R1 B2-8−12.427∗∗0.050∗0.122NS−0.345NS0.016NSPunjab Local × R1 B2-6−3.341NS0.027NS0.098NS0.091NS0.122NSPunjab Local × R1 B2-2910.935∗∗−0.062∗∗−0.150NS−0.150NS−0.143NSPunjab Local × R1 B2-256.611∗∗−0.009NS−0.111NS−0.279NS−0.057NSSE (±)2.0010.0200.1141.1050.135NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 12
Specific combining ability (SCA) effects of mustard hybrids on yield and its attributed traits.
HybridsSUCPRCYPPOYPKranti × R1 B2-23−0.079NS−0.013NS0.305NS−0.043NSKranti × R1 B2-8−0.162NS−0.087NS0.929∗0.353NSKranti × R1 B2-60.131NS0.011NS−0.461NS−0.125NSKranti × R1 B2-29−0.005NS0.055NS−0.670NS−0.256NSKranti × R1 B2-250.115NS0.034NS−0.102NS0.070NSJD 6 × R1 B2-23−0.058NS−0.022NS0.345NS0.048NSJD 6 × R1 B2-8−0.084NS−0.063NS0.715NS0.199NSJD 6 × R1 B2-6−0.114NS−0.022NS0.445NS0.159NSJD 6 × R1 B2-29−0.027NS0.025NS−0.114NS0.066NSJD 6 × R1 B2-250.283NS0.082NS−1.389∗∗−0.472∗R1 B2-5 × R1 B2-230.040NS0.034NS−1.025∗−0.331NSR1 B2-5 × R1 B2-8−0.110NS−0.083NS0.662NS0.299NSR1 B2-5 × R1 B2-6−0.040NS0.018NS−0.065NS−0.036NSR1 B2-5 × R1 B2-29−0.002NS0.028NS−0.147NS−0.147NSR1 B2-5 × R1 B2-250.111NS0.004NS0.575NS0.215NSVaruna × R1 B2-230.119NS−0.064NS0.982∗0.540∗Varuna × R1 B2-80.189NS−0.055NS0.752NS0.245NSVaruna × R1 B2-60.282NS0.043NS−0.631NS−0.227NSVaruna × R1 B2-290.230NS0.010NS−0.067NS−0.078NSVaruna × R1 B2-25−0.820∗∗0.066NS−1.035∗−0.481∗R1 B2-26 × R1 B2-23−0.105NS−0.052NS1.082∗∗0.353NSR1 B2-26 × R1 B2-8−0.055NS−0.016NS−0.215NS−0.165NSR1 B2-26 × R1 B2-6−0.105NS−0.032NS0.402NS0.087NSR1 B2-26 × R1 B2-29−0.001NS0.035NS−0.417NS−0.044NSR1 B2-26 × R1 B2-250.266NS0.065NS−0.852∗−0.231NST 6342 × R1 B2-230.016NS0.028NS−0.661NS−0.256NST 6342 × R1 B2-80.026NS0.034NS−0.591NS−0.237NST 6342 × R1 B2-6−0.081NS−0.015NS0.223NS0.130NST 6342 × R1 B2-29−0.117NS−0.038NS0.814NS0.320NST 6342 × R1 B2-250.156NS−0.008NS0.215NS0.044NSPunjab Local × R1 B2-230.066NS0.090NS−1.028∗−0.311NSPunjab Local × R1 B2-80.196NS0.270NS−2.251∗∗−0.695∗∗Punjab Local × R1 B2-6−0.074NS−0.002NS0.089NS0.012NSPunjab Local × R1 B2-29−0.077NS−0.115NS0.603NS0.139NSPunjab Local × R1 B2-25−0.110NS−0.242NS2.588∗∗0.855∗∗SE (±)0.2650.2390.4100.216NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Punjab Local × R1 B2-25 exhibited a strong unfavorable SCA impact at maturity. This cross is ideal for picking out the early offspring. Cross-combination with high means, correct SCA calculations, and at least one parent with a high GCA, according to Kenga et al. [38], would almost surely improve the desired trait concentration. In this study, Varuna showed a statistically insignificant negative GCA influence on oil yield per crop, but R1 B2-23 had a significant positive GCA effect.The parent Punjab Local had a detrimental effect on plant oil production via GCA. Additional significant GCA effects included siliqua length of the first flowering day to 50% flowering day and maturity days, aphid infestation index, phenol sugar, and protein content, as well as total per plant, primary and secondary branches, siliqua on the main raceme, siliqua on branches, total siliqua per plant, seed number per siliqua, and duration. In this study, significant negative GCA effects on all parameters except maturity time are considered undesirable. GCA has a negative impact on the first flowering day, 50% blooming day, maturity day, aphid infestation index, sugar and protein content, and aphid infestation index. Plant height, number of primary and secondary branches, total siliqua on the main raceme, total siliqua on branches, total number of siliquae per plant, number of seeds per siliqua, weight, maturity duration, harvest index, oil content, and seed yield per plant were all significantly improved by R1 B2-25.Varuna had significant positive GCA effects on oil yield per plant [39], siliqua yield per plant, yield of the top 10 cm of the main raceme, harvest index, and seed yield per crop, whereas Varuna had significant negative GCA effects. These characteristics benefit the breeder. It did, however, exhibit negative GCA effects on undesirable characteristics such as primary and secondary branching, seed number per siliqua, test weight, days to first and 50% flowering, and biological output per plant. The other parent of the cross, R1 B2-23, demonstrated no significant negative GCA effect on oil yield per crop, but a significant negative GCA effect on primary branches, secondary branches, siliqua on main strobili, total number of siliquae, seed number per siliqua, and test weight. As a result, one of the parents in this cross (Varuna) had a high GCA.The Varuna × R1 B2-23 hybrid has a negative effect on plant height because of its SCA content. This cross is perfect for reducing the height of plants. However, for a larger plant, Punjab Local × R1 B2-25 is preferable. Tiwari et al. [40] said that hybrid combinations including at least one good general combiner parent might be used to generate high-producing pure lines via additive gene action, even if SCA effects were negligible.In the present study, the variety Varuna exhibited a negative nonsignificant GCA effect for oil content. It also expressed a negative but nonsignificant general correlating ability effects for earliness and dwarfness, which is in contrary to the earlier findings [39]. This deviation in results may be due to the use of different breeding materials in different environments.Among the biochemical features, no cross exhibited a significant SCA value. Additionally, the aphid infestation index and oil content were similar. Only two hybrids showed a substantial SCA effect on oil production, showing that even if the parents’ grain yields varied widely, they produced hybrids with average oil yields higher than the parents’ limits, although little improvement.


## 3.3. Gene Action, Degree of Dominance, Heritability, Contribution to the Total Variance, and Heterosis

The level of variance attributable to line × testers was greater than that due to testers for the number of seeds per siliqua, biological yield per plant, and stick yield per plant, demonstrating nonadditive variance governing by these traits. To establish the relevance of the GCA and SCA variances produced from the line × tester design, these data appeared exploratory. Therefore, we can calculate the additive (σ2A) and nonadditive (σ2D) components of population variance and the predictability ratio (σ2A/(σ2A+σ2D)). The σ2gca/σ2sca ratio was less than one for all characters, indicating a major involvement of nonadditive gene actions such as dominance, epistasis, and other interaction effects in the expression of these characteristics. Mannuel and Palamisamy [41] and Bobby and Nadarajan [42] found similar results. Because the variance owing to general combining ability (σ2gca) was less than variance due to specific combining ability (σ2sca) for all traits investigated, this supports a predominance of nonadditive gene activity influencing these qualities (Tables 13 and 14). The variation ratio of general to specific combining ability (σ2gca/σ2sca) was less than unity, corroborating these findings. As a result, nonadditive gene results appeared to regulate the heritage of all studied features. Because recombinants within segregating populations might increase features including oil yield, plant height, number of siliquae per plant, and crop length, the selection of superior plants should be deferred to the following generations. In an earlier study using a variety of experimental lines, it was observed that the variance of GCA was greater in plant height and 1000-seed weight from sowing to 50% flowering and maturity, but the variation of SCA was more in seed yield and other remaining boundaries [43]. The discrepancy in the results is primarily attributable to the use of various testing materials. The dominance ratio suggested a predominance of nonadditive gene action in aphid resistance inheritance [44–50]. Negative variance was reported for (σ2D) aphid infestation index, oil content, phenol content, sugar content, and protein content, except for aphid infestation index. Given that they are squared values, the variance cannot be negative. However, they do occur, especially when estimating genetic variance using ANOVA expectation, as in this study to derive σ2gca from ANOVA for combining ability. There are a number of possible explanations for negative variance estimates: (a) small sample size and outlier values; (b) G-x-E interaction that may have increased the error variations; and (c) lack of randomly mating when producing half-sibs. Its validity can be accessed through a series of experiments using an average [51].Table 13
Estimates of genetic variance for yield and its attributed traits in mustard.
ParameterPHPRBSEBSMRSBTOSSDSSLTWDFIFDFFDMMDCovHS (Line)32.2090.3660.6186.9261097.6041293.5490.0070.1020.0055.7676.6540.1154.402CovHS (Tester)9.5831.0060.7315.822421.439531.237−0.0030.0140.0052.4972.6320.0363.078CovHS (Average)2.1290.0680.0670.63977.11892.5540.0000.0060.0000.4190.4710.0080.376CovFS163.8623.5464.23826.7453095.8713694.7680.2710.2840.02715.54017.1790.31614.622Ϭ2gca=2.1290.0680.0670.63977.11892.5540.0000.0060.0000.4190.4710.0080.376Ϭ2sca=92.0780.7231.6382.895437.409484.4070.2670.0940.0090.9400.8900.0560.856Ϭ2gca/Ϭ2sca0.0230.0930.0410.2210.1760.1910.0010.0630.0560.4460.5290.1360.440Ϭ2A=4.2570.1350.1341.279154.236185.1070.0000.0120.0010.8380.9420.0150.753Ϭ2D=92.0780.7231.6382.895437.409484.4070.2670.0940.0090.9400.8900.0560.856Ϭ2A/Ϭ2D=0.0460.1870.0820.4420.3530.3820.0020.1260.1120.8911.0590.2720.880Ϭ2A/(Ϭ2A+Ϭ2D)=0.0440.1570.0760.3060.2610.2760.0020.1120.1010.4710.5140.2140.468h2ns=0.0130.0600.0460.0290.0170.0190.0010.0370.0330.0690.0690.0010.030h2ns (%)=1.3425.9714.6362.8901.7141.9180.0733.6533.2516.9176.9130.1092.995Table 14
Estimates of genetic variance for yield and its attributed traits in mustard.
ParameterYTTBIYSYHIAIIOICPHCSUCPRCYPPOYPCovHS (Line)0.21024.96215.3270.0020.0291.9900.0210.0620.0254.7080.907CovHS (Tester)0.068−5.674−3.9840.0010.0172.4600.0120.0460.0092.2280.671CovHS (Average)0.0141.0170.6000.0000.0020.2220.0020.0050.0020.3510.079CovFS0.52272.44456.3360.0060.0787.6300.0510.1810.01213.2693.005Ϭ2gca=0.0141.0170.6000.0000.0020.2220.0020.0050.0020.3510.079Ϭ2sca=0.04146.11541.2860.002−0.004−0.984−0.008−0.018−0.0480.9260.087Ϭ2gca/Ϭ2sca0.3460.0220.0150.063−0.527−0.225−0.199−0.300−0.0360.3790.916Ϭ2A=0.0282.0341.2010.0000.0050.4430.0030.0110.0040.7020.159Ϭ2D=0.04146.11541.2860.002−0.004−0.984−0.008−0.018−0.0480.9260.087Ϭ2A/Ϭ2D=0.6920.0440.0290.126−1.055−0.451−0.397−0.600−0.0730.7581.831Ϭ2A/(Ϭ2A+Ϭ2D)=0.4090.0420.0280.11219.280−0.821−0.658−1.503−0.0790.4310.647h2ns=0.0580.0190.0160.0410.0320.0690.0330.0350.0130.0530.065h2ns (%)=5.7861.9481.6494.0973.1576.9473.3393.4771.2725.3416.478Furthermore, a predictability ratio greater than 0.5 denotes additive gene activation, whereas one less than 0.5 denotes nonadditive gene measurements. There are a lot of additive and nonadditive gene measurements in a characteristic. Predictability ratios were low and did not surpass 0.5 except for days to 50% flowering, aphid infestation index, and oil production per plant, showing nonadditive gene activity predominates. The prediction ratios for aphid infestation index and oil yield were higher, indicating the dominance of additive gene measures for these variables. This suggests that selection among segregating recombinants can result in plants that are superior in quality in regard to days to 50% blooming, aphid infestation index, and oil yield. All attributes had poor narrow-sense heritability values, according to the present study (Tables13 and 14).Line × testers donated more to the total sum square than any other factors for plant height, secondary branch count, seed count, days to maturity, biological yield, and stick yield (Table15 and Figure 1). Aphid infestation index, phenol and sugar content, seed yield per crop, and oil yield per crop were all determined by lines. In the presence of testers, the variation in primary branch count and oil content was more noticeable. In no case did all three sources of variance contribute equally. Testers and lines together contributed much less to plant height, seed number per siliqua, days to maturity, biological yield, and stick yield per crop. In addition, the interaction between lines and testers influenced the expression of the traits assessed. The current investigation found substantial differences in twenty-four traits between parents and hybrids (Figure 1). The contributions of lines and testers reflect a fixed component of genetic variation, whereas the line × the tester’s element reflects nonfixable factors [52].Table 15
Proportional contribution of lines, testers, and their interactions to total variance.
CharactersProportional contribution of lines, testers, and their interactions to total variance (in %)LinesTestersLines × testersPlant height (cm)32.9414.2852.78No. of primary branches per plant23.9148.0028.09No. of secondary branches per plant29.6328.1042.27No. of siliquae on main raceme38.4029.1732.43No. of siliquae on branches58.9822.5618.46No. of siliquae per plant58.9223.8117.27No. of seeds per siliqua19.4510.7869.77Length of siliqua (cm)49.5311.5738.90Test weight (g)34.3727.7237.92Days to 1st flowering63.6826.389.93Days to 50% flowering64.5124.6210.87Days to maturity32.6214.4452.94Duration of maturity52.6934.4312.88Yield of top 10 cm of the main raceme (g)66.2021.2812.52Biological yield per plant (g)45.771.8152.42Stick yield per plant (g)38.753.6957.57Harvest index44.2418.1537.61Aphid infestation index56.2031.1812.62Oil content (%)44.7451.104.16Phenol content (mg/g)52.6529.0718.28Sugar content (mg/g)44.0930.3525.57Protein content (%)61.2322.6116.16Yield per plant (g)61.3827.7210.91Oil yield per plant(g)55.4438.226.34Figure 1
Proportional contribution of lines, testers, and their interactions to total variance.A remarkable standard heterosis for oil yield per crop was observed in seven hybrids using JD6 as a commercial check: R1 B2-5 × R1 B2-25, Varuna × R1 B2-25, R1 B2-26 × R1 B2-6, and R1 B2-26 × R1 B2-26 (Tables16–20).Table 16
Standard (economic) heterosis of mustard hybrids for yield and its attributed traits.
CrossesPHPRBSEBSMRSBKranti × R1 B2-2315.00∗∗74.31∗∗37.96∗∗−1.22NS88.16∗∗Kranti × R1 B2-818.32∗∗66.15∗∗31.28∗∗−6.44NS86.06∗∗Kranti × R1 B2-624.55∗∗66.73∗∗85.97∗∗−3.18NS89.41∗∗Kranti × R1 B2-2917.36∗∗66.73∗∗32.49∗∗−1.87NS92.64∗∗Kranti × R1 B2-2523.11∗∗65.56∗∗58.62∗∗4.00NS102.71∗∗JD 6 × R1 B2-2318.32∗∗10.76∗22.16∗∗−27.40∗∗106.28∗∗JD 6 × R1 B2-815.41∗∗12.51∗−32.54∗∗−0.89NS90.05∗∗JD 6 × R1 B2-615.18∗∗62.07∗∗67.74∗∗4.65NS101.01∗∗JD 6 × R1 B2-298.94∗67.31∗∗75.64∗∗5.11NS102.36∗∗JD 6 × R1 B2-2520.81∗∗67.90∗∗84.76∗∗4.91NS105.79∗∗R1 B2-5 × R1 B2-2312.86∗∗32.92∗∗20.64∗∗−7.09NS79.59∗∗R1 B2-5 × R1 B2-89.70∗∗20.68∗∗16.69∗−9.04NS74.09∗∗R1 B2-5 × R1 B2-65.86NS31.75∗∗20.34∗∗−4.15NS100.24∗∗R1 B2-5 × R1 B2-2910.51∗∗56.24∗∗27.63∗∗−1.22NS94.46∗∗R1 B2-5 × R1 B2-2513.03∗∗62.65∗∗20.94∗∗4.65NS101.77∗∗Varuna × R1 B2-23−0.45NS35.83∗∗36.14∗∗4.65NS103.26∗∗Varuna × R1 B2-816.04∗∗19.51∗∗34.31∗∗−4.15NS99.90∗∗Varuna × R1 B2-625.15∗∗25.34∗∗37.35∗∗−1.22NS98.78∗∗Varuna × R1 B2-2921.85∗∗45.74∗∗34.92∗∗3.67NS94.87∗∗Varuna × R1 B2-2527.04∗∗64.98∗∗34.31∗∗4.32NS103.57∗∗R1 B2-26 × R1 B2-2326.52∗∗45.74∗∗89.62∗∗6.60NS107.94∗∗R1 B2-26 × R1 B2-824.71∗∗−14.89∗∗0.89NS−3.18NS93.64∗∗R1 B2-26 × R1 B2-625.54∗∗49.82∗∗71.39∗∗4.65NS106.31∗∗R1 B2-26 × R1 B2-2920.02∗∗54.49∗∗51.33∗∗4.65NS114.44∗∗R1 B2-26 × R1 B2-2520.15∗∗69.64∗∗108.46∗∗7.26NS121.62∗∗T 6342 × R1 B2-2326.58∗∗17.18∗∗69.56∗∗−3.83NS91.93∗∗T 6342 × R1 B2-812.20∗∗18.34∗∗52.55∗∗−9.04NS62.85∗∗T 6342 × R1 B2-624.48∗∗23.01∗∗53.15∗∗−0.24NS90.92∗∗T 6342 × R1 B2-2925.92∗∗66.73∗∗58.02∗∗3.87NS106.63∗∗T 6342 × R1 B2-2527.24∗∗66.15∗∗71.99∗∗4.65NS106.11∗∗Punjab Local × R1 B2-2324.55∗∗37.00∗∗−17.35∗−18.82∗∗45.20∗∗Punjab Local × R1 B2-81.88NS−7.89NS−10.05NS−27.63∗∗0.89NSPunjab Local × R1 B2-612.28∗∗21.26∗∗38.57∗∗−14.91∗56.50∗∗Punjab Local × R1 B2-2916.53∗∗26.50∗∗50.72∗∗−9.04NS66.25∗∗Punjab Local × R1 B2-2527.04∗∗54.49∗∗33.71∗∗−1.22NS90.44∗∗NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 17
Standard (economic) heterosis of mustard hybrids for yield and its attributed traits.
CrossesTOSSDSSLTWDFIFKranti × R1 B2-2369.00∗∗−7.52∗∗41.53∗∗6.47∗∗17.43∗∗Kranti × R1 B2-866.24∗∗0.12NS7.00NS−0.70NS24.40∗∗Kranti × R1 B2-669.56∗∗−0.25NS31.20∗∗3.13∗∗20.92∗∗Kranti × R1 B2-2972.39∗∗0.54NS18.80∗∗5.82∗∗16.27∗∗Kranti × R1 B2-2581.55∗∗0.37NS18.80∗∗6.47∗∗12.78∗∗JD 6 × R1 B2-2377.63∗∗−3.32NS5.23NS6.55∗∗12.78∗∗JD 6 × R1 B2-870.56∗∗−5.32∗∗11.79NS2.15∗17.43∗∗JD 6 × R1 B2-680.35∗∗−4.12∗8.26NS1.66NS10.45∗∗JD 6 × R1 B2-2981.51∗∗2.35NS9.54NS2.31∗10.45∗∗JD 6 × R1 B2-2584.17∗∗4.31∗17.37∗5.98∗∗9.29∗∗R1 B2-5 × R1 B2-2361.01∗∗−1.35NS21.12∗∗−8.85∗∗25.57∗∗R1 B2-5 × R1 B2-856.27∗∗0.33NS9.80NS−3.31∗∗26.73∗∗R1 B2-5 × R1 B2-677.86∗∗−4.96∗10.81NS1.34NS20.92∗∗R1 B2-5 × R1 B2-2973.95∗∗−8.41∗∗13.19∗2.15∗19.75∗∗R1 B2-5 × R1 B2-2580.95∗∗−8.15∗∗12.41NS6.72∗∗11.61∗∗Varuna × R1 B2-2382.13∗∗−13.85∗∗12.07NS2.40∗11.61∗∗Varuna × R1 B2-877.60∗∗−7.15∗∗0.65NS0.44NS22.08∗∗Varuna × R1 B2-677.35∗∗−6.37∗∗−5.28NS0.85NS15.10∗∗Varuna × R1 B2-2975.32∗∗0.59NS−3.84NS2.07NS10.45∗∗Varuna × R1 B2-2582.29∗∗−6.32∗∗15.19∗6.80∗∗12.78∗∗R1 B2-26 × R1 B2-2386.22∗∗0.23NS30.22∗∗5.00∗∗9.29∗∗R1 B2-26 × R1 B2-872.89∗∗−10.50∗∗24.27∗∗4.43∗∗22.08∗∗R1 B2-26 × R1 B2-684.52∗∗2.68NS23.68∗∗5.49∗∗11.61∗∗R1 B2-26 × R1 B2-2990.90∗∗−7.00∗∗32.10∗∗5.58∗∗10.45∗∗R1 B2-26 × R1 B2-2597.11∗∗−1.72NS19.61∗∗7.04∗∗9.29∗∗T 6342 × R1 B2-2371.41∗∗−2.23NS22.76∗∗5.17∗∗20.92∗∗T 6342 × R1 B2-847.44∗∗−8.67∗∗1.55NS4.60∗∗26.73∗∗T 6342 × R1 B2-671.38∗∗−2.65NS13.60∗5.33∗∗19.75∗∗T 6342 × R1 B2-2984.60∗∗−4.49∗41.36∗∗5.74∗∗12.78∗∗T 6342 × R1 B2-2584.36∗∗3.63NS18.80∗∗6.96∗∗10.45∗∗Punjab Local × R1 B2-2331.48∗∗−6.06∗∗39.68∗∗2.97∗∗38.36∗∗Punjab Local × R1 B2-8−5.22NS−7.36∗∗35.79∗∗2.56∗55.80∗∗Punjab Local × R1 B2-641.19∗∗−1.19NS28.96∗∗3.86∗∗37.19∗∗Punjab Local × R1 B2-2950.11∗∗−7.10∗∗41.77∗∗4.11∗∗31.38∗∗Punjab Local × R1 B2-2570.79∗∗−5.27∗∗10.18NS6.06∗∗22.08∗∗NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 18
Standard (economic) heterosis of mustard hybrids for yield and its attributed traits.
CrossesDFFDMMDYTTBIYKranti × R1 B2-2311.53∗∗−7.14∗∗−15.45∗∗−5.61NS−8.08NSKranti × R1 B2-817.30∗∗−6.55∗∗−17.02∗∗−13.02∗∗20.10∗∗Kranti × R1 B2-614.41∗∗−6.85∗∗−16.24∗∗−8.96∗13.80∗Kranti × R1 B2-2911.53∗∗−7.14∗∗−15.06∗∗−5.70NS−8.07NSKranti × R1 B2-257.68∗−7.14∗∗−13.88∗∗−22.74∗∗−20.84∗∗JD 6 × R1 B2-237.68∗−7.14∗∗−13.88∗∗3.22NS5.01NSJD 6 × R1 B2-811.53∗∗−7.43∗∗−15.84∗∗−6.23NS1.11NSJD 6 × R1 B2-66.72∗−7.14∗∗−13.09∗∗3.40NS−17.59∗∗JD 6 × R1 B2-292.87NS−6.85∗∗−12.70∗∗5.43NS17.48∗∗JD 6 × R1 B2-251.91NS−7.43∗∗−13.09∗∗−21.59∗∗−8.73NSR1 B2-5 × R1 B2-2315.37∗∗−7.43∗∗−18.60∗∗−16.20∗∗−22.57∗∗R1 B2-5 × R1 B2-816.33∗∗−8.02∗∗−19.78∗∗−18.76∗∗−27.64∗∗R1 B2-5 × R1 B2-611.53∗∗−7.43∗∗−17.02∗∗−10.73∗∗−26.70∗∗R1 B2-5 × R1 B2-2910.57∗∗−7.43∗∗−16.63∗∗−6.23NS−22.92∗∗R1 B2-5 × R1 B2-253.84NS−7.43∗∗−13.88∗∗−22.65∗∗−22.68∗∗Varuna × R1 B2-233.84NS−7.43∗∗−13.88∗∗3.93NS−25.99∗∗Varuna × R1 B2-812.49∗∗−8.32∗∗−18.60∗∗−8.61∗−21.79∗∗Varuna × R1 B2-66.72∗−7.73∗∗−15.45∗∗−5.08NS−27.61∗∗Varuna × R1 B2-292.87NS−6.85∗∗−12.70∗∗0.22NS−17.09∗∗Varuna × R1 B2-254.80NS−7.14∗∗−13.88∗∗−22.47∗∗−14.71∗R1 B2-26 × R1 B2-231.91NS−6.85∗∗−12.30∗∗4.81NS2.69NSR1 B2-26 × R1 B2-818.26∗∗−7.14∗∗−17.02∗∗−7.99∗−23.15∗∗R1 B2-26 × R1 B2-69.60∗∗−7.73∗∗−14.27∗∗3.40NS−0.52NSR1 B2-26 × R1 B2-298.64∗∗−7.14∗∗−13.09∗∗4.28NS−12.91∗R1 B2-26 × R1 B2-257.68∗−7.14∗∗−12.70∗∗5.08NS−5.73NST 6342 × R1 B2-2317.30∗∗−7.73∗∗−17.42∗∗−9.23∗3.61NST 6342 × R1 B2-822.10∗∗−7.73∗∗−19.38∗∗−21.06∗∗12.00∗T 6342 × R1 B2-616.33∗∗−6.85∗∗−15.84∗∗−5.61NS−2.60NST 6342 × R1 B2-2910.57∗∗−5.67∗∗−11.91∗∗2.52NS−13.70∗T 6342 × R1 B2-258.64∗∗−6.26∗∗−11.91∗∗−21.06∗∗−9.12NSPunjab Local × R1 B2-2331.72∗∗−7.73∗∗−23.32∗∗−37.57∗∗−31.12∗∗Punjab Local × R1 B2-843.26∗∗−5.97∗∗−26.86∗∗−55.50∗∗−57.74∗∗Punjab Local × R1 B2-627.87∗∗−5.67∗∗−20.17∗∗−32.45∗∗−34.80∗∗Punjab Local × R1 B2-2923.07∗∗−5.67∗∗−18.20∗∗−24.33∗∗0.28NSPunjab Local × R1 B2-2515.37∗∗−7.14∗∗−17.02∗∗−34.13∗∗−9.26NSNS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 19
Standard (economic) heterosis of mustard hybrids for yield and its attributed traits.
CrossesSYHIAIIOICPHCKranti × R1 B2-23−14.87NS13.70NS−29.12∗∗−7.63NS−8.29∗Kranti × R1 B2-830.03∗∗−21.14∗−22.37∗∗−10.24∗−4.84NSKranti × R1 B2-619.04∗−13.00NS−25.74∗∗−2.67NS−6.57∗Kranti × R1 B2-29−14.59NS13.02NS−27.99∗∗−1.66NS−7.71∗Kranti × R1 B2-25−37.90∗∗45.19∗∗−33.62∗∗1.46NS−10.59∗∗JD 6 × R1 B2-23−0.73NS9.44NS−33.62∗∗−4.85NS−10.59∗∗JD 6 × R1 B2-8−1.21NS2.37NS−30.24∗∗−9.32∗−8.86∗∗JD 6 × R1 B2-6−33.22∗∗39.44∗∗−32.49∗∗−2.33NS−10.01∗∗JD 6 × R1 B2-2915.98NS0.09NS−31.37∗∗0.95NS−9.44∗∗JD 6 × R1 B2-25−21.37∗∗28.11∗∗−33.62∗∗1.97NS−7.83∗R1 B2-5 × R1 B2-23−30.26∗∗19.34∗−22.37∗∗−0.22NS−4.84NSR1 B2-5 × R1 B2-8−36.18∗∗23.56∗−22.37∗∗−5.53NS−4.84NSR1 B2-5 × R1 B2-6−38.66∗∗33.64∗∗−25.74∗∗0.20NS−6.57∗R1 B2-5 × R1 B2-29−35.80∗∗34.42∗∗−29.12∗∗0.62NS−8.29∗R1 B2-5 × R1 B2-25−40.86∗∗49.59∗∗−33.62∗∗3.14NS−9.44∗∗Varuna × R1 B2-23−45.64∗∗56.32∗∗−32.49∗∗−3.00NS−10.70Varuna × R1 B2-8−32.74∗∗28.42∗∗−25.74∗∗−10.83∗∗−6.57∗Varuna × R1 B2-6−43.06∗∗44.76∗∗−29.12∗∗−3.59NS−8.29∗Varuna × R1 B2-29−32.36∗∗38.26∗∗−32.49∗∗−2.83NS−10.13∗∗Varuna × R1 B2-25−29.49∗∗35.87∗∗−35.87∗∗−1.57NS−11.74∗∗R1 B2-26 × R1 B2-23−6.37NS16.88NS−36.99∗∗2.72NS−12.43∗∗R1 B2-26 × R1 B2-8−33.60∗∗27.52∗∗−25.74∗∗−0.98NS−6.57∗R1 B2-26 × R1 B2-6−8.76NS15.70NS−35.87∗∗4.41NS−13.00∗∗R1 B2-26 × R1 B2-29−27.01∗∗33.28∗∗−35.87∗∗8.53∗−11.74∗∗R1 B2-26 × R1 B2-25−18.79∗28.09∗∗−36.99∗∗9.46∗−13.58∗∗T 6342 × R1 B2-234.33NS−4.25NS−25.74∗∗−5.87NS−6.57∗T 6342 × R1 B2-822.39∗∗−23.35∗−18.99∗∗−11.76∗∗−3.12NST 6342 × R1 B2-6−6.85NS6.95NS−31.37∗∗−3.34NS−9.44∗∗T 6342 × R1 B2-29−27.29∗∗32.30∗∗−32.49∗∗−2.16NS−8.98∗∗T 6342 × R1 B2-25−22.33∗∗29.61∗∗−38.12∗∗−1.15NS−12.89∗∗Punjab Local × R1 B2-23−30.45∗∗−4.89NS−14.49∗∗−4.52NS−2.08NSPunjab Local × R1 B2-8−59.01∗∗4.12NS−5.49NS−12.52∗∗0.10NSPunjab Local × R1 B2-6−38.66∗∗10.57NS−12.24∗−4.34NS−0.93NSPunjab Local × R1 B2-297.48NS−18.60∗−23.49∗∗−3.26NS−6.68∗Punjab Local × R1 B2-25−15.26NS11.97NS−25.74∗∗−1.99NS−6.57∗NS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Table 20
Standard (economic) heterosis of mustard hybrids for yield and its attributed traits.
CrossesSUCPRCYPPOYPKranti × R1 B2-23−1.34NS−2.19NS0.85NS−6.90NSKranti × R1 B2-80.20NS−1.84NS−8.60∗−18.02∗∗Kranti × R1 B2-61.31NS−2.07NS−4.46NS−7.06NSKranti × R1 B2-29−1.45NS−2.16NS0.25NS−1.45NSKranti × R1 B2-25−3.51NS−2.64NS10.92∗∗12.47∗JD 6 × R1 B2-23−3.51NS−2.66NS10.88∗∗5.44NSJD 6 × R1 B2-8−1.38NS−2.14NS−0.14NS−9.49∗JD 6 × R1 B2-6−3.65NS−2.66NS10.90∗∗8.26NSJD 6 × R1 B2-29−4.07NS−2.72NS13.46∗∗14.48∗∗JD 6 × R1 B2-25−4.14NS−2.83NS12.78∗∗14.96∗∗R1 B2-5 × R1 B2-230.09NS−1.82NS−10.84∗∗−11.08∗R1 B2-5 × R1 B2-80.93NS−1.69NS−13.75∗∗−18.57∗∗R1 B2-5 × R1 B2-6−0.29NS−1.91NS−5.53NS−5.39NSR1 B2-5 × R1 B2-29−1.24NS−2.16NS−0.03NS0.53NSR1 B2-5 × R1 B2-25−3.37NS−2.66NS11.57∗∗15.02∗∗Varuna × R1 B2-23−3.47NS−2.67NS11.61∗∗8.22NSVaruna × R1 B2-8−0.33NS−1.91NS−3.09NS−13.63∗∗Varuna × R1 B2-6−1.31NS−2.14NS1.09NS−2.59NSVaruna × R1 B2-29−3.19NS−2.61NS10.57∗∗7.38NSVaruna × R1 B2-25−17.52∗∗−2.71NS11.78∗∗9.97∗R1 B2-26 × R1 B2-23−3.86NS−2.79NS15.82∗∗18.92∗∗R1 B2-26 × R1 B2-8−0.92NS−1.89NS−5.45NS−6.41NSR1 B2-26 × R1 B2-6−3.40NS−2.69NS11.04∗∗15.88∗∗R1 B2-26 × R1 B2-29−3.65NS−2.66NS12.00∗∗21.49∗∗R1 B2-26 × R1 B2-25−4.17NS−2.89NS16.49∗∗27.45∗∗T 6342 × R1 B2-23−0.78NS−1.89NS−4.32NS−9.97∗T 6342 × R1 B2-81.73NS−1.14NS−17.18∗∗−26.96∗∗T 6342 × R1 B2-6−1.34NS−2.11NS0.52NS−2.88NST 6342 × R1 B2-29−3.05NS−2.52NS10.16∗∗7.72NST 6342 × R1 B2-25−3.51NS−2.76NS13.64∗∗12.29∗Punjab Local × R1 B2-234.84NS0.28NS−36.80∗∗−39.69∗∗Punjab Local × R1 B2-88.61∗1.90NS−57.61∗∗−62.93∗∗Punjab Local × R1 B2-63.83NS−0.18NS−30.53∗∗−33.58∗∗Punjab Local × R1 B2-292.47NS−1.05NS−21.36∗∗−23.95∗∗Punjab Local × R1 B2-25−1.20NS−2.07NS−1.98NS−3.97NSNS: nonsignificant;∗ and ∗∗: significant effects at 5% and 1% level of significance, respectively.Aside from oil yield heterosis, all 10 crosses showed significant plant height heterosis. R1 B2-26 × R1 B2-25 exhibited the highest oil yield per plant heterosis and a positive plant height heterosis. The total number of primary branches, secondary branches, siliquae on branches, and total siliquae per plant and siliqua length are all factors to consider. The day to first flowering and day to 50% flowering of a plant are used to calculate its seed yield. Days to maturity, stick production per plant, aphid, infestation index, phenol, sugar, and protein contents are all negative heterosis in this cross (Tables16–20). For most of the major characters, including seed yield, Ghosh et al. [53] found that both additive and nonadditive gene actions were critical. Plant height and yield are affected by nonadditive gene action, according to Premlatha et al. [54]. On seed weight and plant height, Gnanasekaran et al. [55] discovered a nonadditive gene effect, whereas Sharma [56] discovered an additive gene effect. Verma et al. [57] found σ2sca variation over σ2gca in barley. Both Borghi et al. [58] and Borghi and Perenzin [59] discovered that 2gca was more essential than σ2sca for most characters. Nonadditive σ2sca is most common in space planting, says Lucken [60]. The disparity in results between research projects may be due to differences in breeding material and genotype conditions.


## 4. Conclusion

The mean oil yield of R1 B2-26 × R1 B2-25, JD6 × R1 B2-25, and JD6 × R1 B2-29 hybrids was better than that of the parents. The variance accounted for by SCA was greater than that accounted for by GCA, as indicated by theσ2gca/σ2sca ratio being less than one for all characters, implying a significant role for nonadditive gene actions such as dominance, epistasis, and other interaction effects in the appearance of these traits. The line × tester and lines both played a significant role in the variation in the expression of the majority of the traits studied. Because of the testers, there was more variation in expression for total primary branches per plant, oil content, and oil production per plant. All three sources of variance were not equally significant in any case. The best general combiners for oil yield per plant among the lines are JD6 and Varuna, while the best general combiner among the testers is R1 B2-25. JD6 is an excellent combiner for reducing plant height. Punjab Local is the best all-purpose combiner for shortening the crop’s duration. Punjab Local × R1 B2-25 and Varuna × R1 B2-23 are the two best specific crosses for increasing oil yield, while Varuna × R1 B2-23 is the best for reducing plant height. Three of the finest cross combinations for oil yield per plant are R1 B2-26 × R1 B2-25, R1 B2-26 × R1 B2-29, and R1 B2-26 × R1 B2-23. Additionally, the cross R1 B2-26 × R1 B2-25 exhibited positive heterosis for total siliqua per plant and test weight but negative heterosis for the aphid infestation index. The predominance of nonadditive gene actions suggested unequivocally that the selection of superior individuals should be deferred to subsequent generations.



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*Source: 4387318-2022-05-24.xml*