Determination of general combining ability and Heterosis for yield and quality of tomato inbred lines

doi:10.21203/rs.3.rs-5229104/v1

This study investigates the genetic potential of various lines and testers for breeding purposes, emphasizing the importance of General Combining Ability (GCA) and Specific Combining Ability (SCA) in enhancing crop traits. Lines NBLTM-3, NBLTM-4, and NBLTM-16, along with testers NBLTM-21, NBLTM-24, NBLTM-26, and NBLTM-28, exhibited high GCA, indicating their consistent positive contributions to trait performance. Notably, 68% of the evaluated hybrids displayed high SCA across fourteen traits, highlighting effective combinations and the predominance of non-additive gene action. The analysis revealed that line contributions were more significant than those of testers, with line × tester interactions playing a critical role in hybrid performance. Twelve hybrids demonstrated high overall heterotic status, with NBLTM-4 × NBLTM-29 and NBLTM-3 × NBLTM-28 leading in heterotic responses. Positive heterosis was observed in key traits such as fruit weight and total soluble solids, while negative heterosis was noted in traits like blossom end size. These findings underscore the potential for developing superior hybrids to boost agricultural productivity by focusing on the strengths of both parental lines and hybrid selection.

Horticulture

Virology

Population Genetics

Tomato genotypes

ToLCV

Ty gene

Tester

Lines

General Combining Ability

Specific Combining Ability

Heterosis

Tomato (Solanum lycopersicum L.) is one of the most economically significant vegetable crops worldwide, valued for its nutritional properties and culinary versatility (Salunkhe et al., 1987). As global population increases, ensuring food security necessitates the development of high-yielding tomato varieties with enhanced quality traits such as flavor, firmness, and nutritional content (Pruthi, 1993). Achieving these objectives requires a comprehensive understanding of the genetic factors governing yield and quality attributes in tomato.

Genetic improvement in tomatoes often involves hybridization, wherein desirable traits from different parental lines are combined to generate superior hybrid varieties. The success of hybrid breeding programs relies on the identification and utilization of inbred lines possessing favorable genetic attributes (Metwally et al., 2015). Assessing the general combining ability (GCA) of these inbred lines provides insights into their overall breeding value, particularly their additive genetic effects contributing to trait expression in hybrids (Vijeth et al., 2018).

Heterosis, commonly referred to as hybrid vigor, describes the phenomenon where hybrid offspring exhibit superior performance relative to their parents for certain traits (Sridhara et al., 2008). Heterosis plays a crucial role in plant breeding, as it offers opportunities to develop hybrids with enhanced yield and quality characteristics compared to their parental lines. Understanding the extent and nature of heterosis in tomato hybrids is essential for effective hybrid variety development and deployment (Singh and Asati, 2011).

This study aims to determine the GCA, SCA and heterosis for yield and quality traits in a set of tomato inbred lines. By conducting controlled hybridization experiments and rigorous statistical analyses, we seek to identify inbred lines with high GCA for targeted trait improvement and evaluate the potential for exploiting heterosis in hybrid tomato breeding. The findings of this research will contribute to the development of superior tomato varieties with increased yield and improved quality, addressing the evolving demands of consumers and the agricultural industry.

1. Selection of Inbred Lines : Identify a diverse set of tomato inbred lines known for their distinct phenotypic traits related to yield and quality. Choose lines with contrasting characteristics to maximize genetic diversity within the breeding population.

2. Experimental Design: Establish field trial plots in a randomized complete block design (RCBD) to minimize experimental error and environmental variability. Replicate each hybrid and parental line across multiple blocks to account for within-field variability. Allocate buffer rows between different genotypes to minimize cross-pollination.

3. Field Trials : Prepare the field site by ensuring proper soil preparation, irrigation, and weed management. Plant seeds or seedlings of parental lines and hybrids at appropriate spacing and row arrangements. Implement standard agronomic practices throughout the growing season, including fertilization, pest control, and irrigation.

5. Trait Measurement: Record data on yield-related traits, including fruit weight, number of fruits per plant, total yield per plant and fruit size. Assess quality traits such as fruit firmness, color, sugar content (Brix), carotenoids. Utilize standardized measurement techniques and equipment to ensure accuracy and consistency across observations.

5. Statistical Analysis: Perform analysis of variance (ANOVA) to partition the total variation into components attributable to genotypes (inbred lines and hybrids) and environments. Estimate general combining ability (GCA) effects for each parental line using appropriate statistical methods, such as Griffing's method or analysis of variance of specific combining ability (ANOVASCA). Calculate specific combining ability (SCA) effects for hybrid combinations to assess non-additive genetic effects. Determine heterosis for yield and quality traits by comparing the performance of hybrids to their mid-parent values (MPVs) and parental lines.

6. Validation and Replication: Validate the findings through repeated field trials across multiple growing seasons and environments. Replicate the experiment to confirm the stability and consistency of GCA, SCA, and heterosis estimates.

Mean performance of parents and hybrids for yield and yield attributes

Mean performance results of parents (Lines and Testers) and hybrids for 14 quantitative traits are presented (table 1 a, 1 b, 1c, 1 d, 1 e and 1 f) and discussion are presented.

1. Lines:

The line NBLTM-23 recorded the highest fruit weight (81.36 g), while NBLTM-16 had the lowest (37.8 g). For total soluble sugars, NBLTM-23 exhibited the highest Brix value (5.10), and NBLTM-4 and NBLTM-17 had the lowest (4.15). In terms of fruit length, NBLTM-17 showed the greatest length (6.45 cm), whereas NBLTM-23 had the shortest (4.15 cm). NBLTM-16 also had the highest fruit width (6.55 cm), with NBLTM-3 being the narrowest (4.95 cm). The thickness of the pericarp was highest in NBLTM-3 (0.775 mm) and lowest in NBLTM-23 (0.575 mm). The largest size of locules was observed in NBLTM-3 (3.85 cm), while NBLTM-16 had the smallest (1.65 cm). NBLTM-4 recorded the highest fruit firmness (10.86 kg/cm²), in contrast to NBLTM-16, which had the lowest (2.73 kg/cm²). For fruit size at the blossom end, NBLTM-3 was the largest (0.33 mm), and NBLTM-4 was the smallest (0.13 mm). The highest number of locules was found in NBLTM-16 (5), while NBLTM-3 had the lowest (2). The length of the fruit stalk was greatest in NBLTM-3 (3.45 cm) and shortest in NBLTM-4 (1.90 cm). NBLTM-16 produced the highest seed yield per fruit (323.5 mg), whereas NBLTM-3 had the lowest (130.0 mg). Additionally, NBLTM-3 had the highest number of seeds per fruit (76.5), while NBLTM-17 had the lowest (34.0). NBLTM-23 showed the best seed germination (79.5%), with NBLTM-16 recording the lowest (37.5%). The electrical conductivity was lowest in NBLTM-17 (31.79 μS/cm/g) and highest in NBLTM-4 (68.81 μS/cm/g) as exhibited in table 1 a and 1 b.

2. Testers:

Among the testers, NBLTM-28 had the highest fruit weight (100.05 g), while NBLTM-24 recorded the lowest (50.55 g). For total soluble sugars, NBLTM-28 also led with a Brix value of 6.20, and NBLTM-24 had the lowest (4.45). The highest fruit length was observed in NBLTM-28 (6.25 cm), compared to the lowest in NBLTM-21 (4.25 cm). NBLTM-28 also had the widest fruit (5.55 cm), while NBLTM-24 had the narrowest (4.45 cm). In thickness of pericarp, NBLTM-28 again ranked highest (0.88 mm) and NBLTM-21 the lowest (0.55 mm). The largest size of locules was found in NBLTM-28 (2.75 cm), and NBLTM-26 had the smallest (2.25 cm). The highest firmness was in NBLTM-21 (8.85 kg/cm²), while NBLTM-26 had the lowest (5.18 kg/cm²). For fruit size at the blossom end, NBLTM-28 was the largest (0.35 mm), and NBLTM-24 was the smallest (0.19 mm). NBLTM-21 had the highest number of locules (6), whereas NBLTM-28 had the lowest (2). The length of the fruit stalk was highest in NBLTM-28 (2.90 cm) and lowest in NBLTM-29 (2.40 cm). NBLTM-21 recorded the highest seed yield per fruit (239.5 mg), while NBLTM-28 had the lowest (144.0 mg). The highest number of seeds per fruit was in NBLTM-21 (63.0), compared to the lowest in NBLTM-26 (33.5). NBLTM-28 exhibited the highest seed germination (94.5%), while NBLTM-24 had the lowest (47.5%). The electrical conductivity was lowest in NBLTM-28 (33.60 μS/cm/g) and highest in NBLTM-29 (69.38 μS/cm/g) as exhibited in table 1 c and 1 d.

3. Hybrids (Parental Crosses):

Among the parental crosses, NBLTM-16 × NBLTM-26 had the highest fruit weight (138.5 g), while NBLTM-23 × NBLTM-28 recorded the lowest (38.4 g). For total soluble sugars, the highest was in NBLTM-4 × NBLTM-21 (6.25 Brix), and the lowest was observed in NBLTM-4 × NBLTM-29 (3.35 Brix). The hybrid NBLTM-16 × NBLTM-28 had the longest fruit (6.62 cm), while NBLTM-23 × NBLTM-28 had the shortest (2.82 cm). In terms of fruit width, NBLTM-16 × NBLTM-26 was the widest (7.89 cm), and NBLTM-17 × NBLTM-29 was the narrowest (2.30 cm). The thickness of pericarp was highest in NBLTM-4 × NBLTM-28 (1.25 mm), while the lowest was in NBLTM-17 × NBLTM-24 and NBLTM-23 × NBLTM-28 (0.36 mm). The largest size of locules was in NBLTM-4 × NBLTM-26 (4.05 cm), and the smallest was in NBLTM-17 × NBLTM-29 (1.15 cm). The highest firmness among hybrids was observed in NBLTM-3 × NBLTM-24 (11.80 kg/cm²), and the lowest in NBLTM-3 × NBLTM-28 (5.31 kg/cm²). For fruit size at the blossom end, the highest was recorded in NBLTM-3 × NBLTM-24, NBLTM-3 × NBLTM-29, and NBLTM-16 × NBLTM-28 (0.19 mm), while the lowest was found in NBLTM-3 × NBLTM-28, NBLTM-4 × NBLTM-21, and NBLTM-4 × NBLTM-24 (0.09 mm). The highest number of locules was in NBLTM-16 × NBLTM-26 (6). Lastly, the length of the fruit stalk was greatest in NBLTM-3 × NBLTM-21 (3.20 cm) and lowest in NBLTM-4 × NBLTM-29 (2.25 cm). The highest seed yield per fruit among hybrids was in NBLTM-4 × NBLTM-28 (390.0 mg), while the lowest was in NBLTM-17 × NBLTM-29 (80.0 mg). The highest number of seeds per fruit was in NBLTM-16 × NBLTM-26 (166.5), while the lowest was in NBLTM-4 × NBLTM-29 (24.0). The highest seed germination was recorded in NBLTM-17 × NBLTM-21 (88.5%), and the lowest was in NBLTM-16 × NBLTM-26 and NBLTM-23 × NBLTM-26 (70.5%). The lowest electrical conductivity was in NBLTM-17 × NBLTM-21 (18.42 μS/cm/g), while the highest was in NBLTM-3 × NBLTM-28 (63.58 μS/cm/g) as exhibited in table 1 e and 1 f.

Analysis of variance (ANOVA)

The analysis of variance due to different sources of variation for fourteen traits studied viz., weight of fruits, total soluble solids, fruit length, fruit width, thickness of pericarp, size of locules, fruit firmness, size at blossom end, number of locules, length of fruit stalk, seed yield per fruit, number of seeds per fruit, seed germination and electrical conductivity are summarized in (Table 2 a and 2 b).

The ANOVA indicated that, the difference among the genotypes (parents and crosses) and variance due to crosses was significant for all the characters based on their mean sum of square values. The difference among the parents was highly significant for all the traits. The variances due to lines and testers were significant variance for all the traits studied and also the line × tester interaction showed variance was significant for all the traits. Signiﬁcant variances due to line × tester interaction for all the characters suggested the presence of signiﬁcant variances for sca among hybrids (Vekariya et al., 2019). The results on the mean performance of parents for fourteen traits studied revealed substantial variability among lines and testers. However, in general no single line or tester was found to have better mean performance for all the characters studied. These results are in agreement with those of Metwally et al. (2015), Nayana et al. (2021) and Navaneet Singh et al. (2022).

Combining ability analysis

The general combining ability and specific combining ability effects of parents (lines and testers) estimated for fourteen traits is presented in (Table 3 a, 3 b, 3c, 3 d and 5 a and 5 b).

The combining ability analysis revealed significant effects across various traits. For fruit weight, NBLTM-4 and NBLTM-16 showed positive GCA, while NBLTM-17 and NBLTM-23 had negative effects; the hybrid NBLTM-23 × NBLTM-29 exhibited the highest positive SCA. In terms of total soluble sugars, NBLTM-4 and NBLTM-16 demonstrated positive GCA, with NBLTM-4 × NBLTM-24 showing the highest positive SCA. For fruit length, positive GCA was observed in NBLTM-3 and NBLTM-16, while the hybrid NBLTM-17 × NBLTM-21 had the highest positive SCA. Regarding fruit width, NBLTM-16 and NBLTM-4 were positive for GCA, with NBLTM-3 × NBLTM-29 displaying the highest positive SCA. In thickness of pericarp, NBLTM-3 and NBLTM-4 had positive GCA, and NBLTM-17 × NBLTM-21 showed the highest positive SCA. For size of locules, NBLTM-3 and NBLTM-4 exhibited positive GCA, with NBLTM-17 × NBLTM-21 having the highest positive SCA. Fruit firmness was positive in NBLTM-17 but negative in NBLTM-23; NBLTM-3 × NBLTM-24 had the highest positive SCA. Seed yield per fruit showed a positive GCA in NBLTM-16, while all hybrids had non-significant SCA effects. In number of seeds per fruit, NBLTM-16 had a positive GCA and NBLTM-29 a negative one, with all hybrids showing non-significant SCA effects. For seed germination, NBLTM-3, NBLTM-4, and NBLTM-17 showed positive GCA, while NBLTM-16 and NBLTM-23 were negative; the hybrid NBLTM-16 × NBLTM-28 had the highest positive SCA. Lastly, in electrical conductivity, NBLTM-3 and NBLTM-4 had positive GCA, while the hybrid NBLTM-3 × NBLTM-24 exhibited the highest negative SCA. This trait-wise analysis highlights the diverse combining abilities of lines, testers, and hybrids, offering insights for future breeding efforts.

The combining ability analysis revealed significant differences among the lines, testers, and hybrids for various traits. In terms of General Combining Ability (GCA), lines such as NBLTM-3, NBLTM-4, and NBLTM-16 exhibited high positive GCA for several traits, including fruit weight, total soluble sugars, and fruit length, while lines like NBLTM-17 and NBLTM-23 showed consistently negative GCA across multiple traits. Among the testers, NBLTM-21, NBLTM-24, NBLTM-26, and NBLTM-28 demonstrated positive GCA, particularly in fruit width and seed yield, whereas NBLTM-29 often yielded negative GCA. For Specific Combining Ability (SCA), notable hybrids like NBLTM-23 × NBLTM-29 and NBLTM-3 × NBLTM-21 showed significant positive effects for fruit weight and total soluble sugars, while hybrids such as NBLTM-16 × NBLTM-21 exhibited negative SCA effects in several traits. Overall, the analysis indicates that specific lines and hybrids hold promise for enhancing desirable characteristics in future breeding programs.

Overall general combining ability status of lines and testers

It can be substantiated from the (Table 4 a and 4 b) that the lines NBLTM-3, NBLTM-4 and NBLTM-16 were found to possess high (H) overall general combining ability status based on their rankings. However, the lines NBLTM-17 and NBLTM-23 were found to possess low (L) overall general combining ability effects based on their respective rankings. Out of five testers, four testers viz., NBLTM-21, NBLTM-24, NBLTM-26 and NBLTM-28 recorded high (H) overall general combining ability status and NBLTM-29 was found to possess low (L) overall general combining ability effects based on their respective rankings (Table 4 a and 4 b).

The selection of potential parent in hybridization may be judged by comparing the per se performance of the parents, the F₁value (heterosis) and the combining ability effects. However, per se performance of a parent alone is not always a true indicator of its potentiality in hybrid combination most of the time. Therefore, general combining ability, which is also the breeding value of the parent, has proved as a useful tool for selecting the parents for hybridization. Among parents, the significant gca effects was observed, the parents with higher magnitude of gca effects were considered as superior to those with lower magnitude. Results revealed that 80.0 % of the testers and 60.0 % of the lines performed better and have overall general combining ability in an estimated direction. The superiority of female and male lines were observed for gca. These results are on par with the reports of Zengin et al. (2015); Venkariya et al. (2019); Liu et al. (2021) and Navneet Singh et al. (2022).

There is a need of identifying the superior combiners which helps the breeders in selecting appropriate parents to be used in the breeding programmes to develop superior hybrids in tomato (Liu et al., 2021).

Overall specific combining ability status of hybrids

It is important to determine whether hybrids possess higher sca values across all the characters or not. The overall sca status of all the hybrids across fourteen traits was determined and same is presented in (Table 6). From the results, it is evident that 68 % of the hybrids, i.e., 17 of the twenty five hybrids had high (H) overall sca status and the remaining eight had low (L) overall sca status across the fourteen traits.

Several workers including Vekariya et al. (2019); Nayana et al. (2021) and Navneet Singh et al. (2022) identified good specific combiners for different yield and yield attributing characters based on high sca effects in desirable direction. One of the parent with high gca another parent with low gca, resulted in hybrid with high sca effect, which may be due to the predominance of dominant allele Vekariya et al. (2019). When both the parents recorded low gca effects, led to a hybrid with high sca effect which may be due to accumulation of favourable alleles and non-additive gene action. Similar findings as observed in the present study were also reported by Singh and Asati (2011); Zengin et al. (2015); Metwally et al. (2015); Vekariya et al. (2019); Nayana et al. (2021) and Navneet Singh et al. (2022) in tomato.

Variance due to general and specific combining ability effects

The variance due to general and specific combining ability effects for fourteen different traits is presented in (Table 7, Fig. 1). The results revealed that, an estimate of sca variance was predominant for all the characters studied as indicated by gca to sca ratio. This was maximum in respect of seed yield per fruit (1885.275) followed by weight of fruit (434.456) and number of seeds per fruit (276.578). The predominance of sca variance indicated there is a presence of non-additive gene action.

These results indicated that preponderance of non-additive gene action in the expression of all the traits studied and a very good prospect for the exploitation of non-additive genetic variation for traits through hybrid breeding in tomato (Zengin et al. 2015). The estimates of variance due to gca and sca indicated the predominance of sca variance for all the traits. These results are in conformity with Singh and Asati (2011); Zengin et al. (2015); Metwally et al. (2015); Vekariya et al. (2019); Nayana et al. (2021) and Navneet Singh et al. (2022) in tomato and also reported that non-additive gene effects were predominant than additive gene effects especially for yield and its component characters. The presence of non-additive genetic variance in this study offer scope for exploitation of heterosis for these characters.

Proportional contribution of lines, testers and line × tester interaction towards the performance of hybrids

The proportional contribution of lines, testers and lines × testers interaction for nine traits is presented in (Table 8 and Fig. 2).

The results revealed that the per cent contribution of lines was higher for all the traits compared with the testers. However, the per cent contribution of lines × testers interaction was more for all the characters.

The results revealed that contribution per cent contribution of the line × tester interaction effect to the variation observed in hybrids was higher for all the traits. The lines contributed towards the variation in the hybrids for all the traits. Overall lines were found to have more role in the variation among the hybrids. Therefore, emphasis needs to be given while selecting the lines. However, the same thing cannot be done with respect to lines × testers interaction as it is not under the breeder's control, but it depends on the particular way in which the lines and testers interact and also on the environment in which the hybrids are grown. The results were on par with Navneet Singh et al. (2022) in tomato.

Heterosis

The results of heterosis of fourteen quantitative traits for important yield attributing traits are presented and discussed hereunder (Table 9 a, b, c, d and e).

Mid Parent Heterosis

Across the evaluated traits, significant positive heterosis over the mid parent was observed in several hybrids. Notably, fruit weight exhibited impressive gains, particularly with NBLTM-16 × NBLTM-24 showing 160.78%. In terms of total soluble solids, NBLTM-4 × NBLTM-24 led at 39.73%. Fruit length also demonstrated positive heterosis, with NBLTM-16 × NBLTM-26 achieving 42.10%. Other traits, such as fruit width (NBLTM-16 × NBLTM-26 at 31.58%) and thickness of pericarp (NBLTM-4 × NBLTM-26 at 91.67%), followed suit. For seed germination, hybrids like NBLTM-16 × NBLTM-24 exhibited 68.24%, while fruit firmness peaked with NBLTM-16 × NBLTM-26 at 104.30%. Conversely, several traits, including size at blossom end and number of locules, revealed significant negative heterosis, with NBLTM-3 × NBLTM-28 showing -80.05% and NBLTM-4 × NBLTM-21 at -60.00%, respectively.

Better Parent Heterosis

When comparing hybrids to the better parent, several traits exhibited substantial positive heterosis. For fruit weight, NBLTM-16 × NBLTM-24 achieved 132.09%, and total soluble solids were notably enhanced by NBLTM-4 × NBLTM-24 at 34.83%. In fruit length, NBLTM-16 × NBLTM-26 reached 41.32%, while fruit width showed NBLTM-16 × NBLTM-26 at 20.53%. The thickness of pericarp also demonstrated improvements with NBLTM-4 × NBLTM-26 at 58.62%. However, for certain traits like size of locules and number of locules, significant negative heterosis was noted against the better parent, with NBLTM-4 × NBLTM-21 reaching -66.67%. For seed yield per fruit and number of seeds per fruit, improvements were limited, with the best results being NBLTM-4 × NBLTM-28 and NBLTM-16 × NBLTM-24 showing positive heterosis of 109.40% and 180.90%, respectively.

Standard Heterosis

In terms of standard heterosis, several hybrids demonstrated remarkable improvements across various traits. For fruit weight, the hybrid NBLTM-16 × NBLTM-26 exhibited a striking 265.92%, while NBLTM-16 × NBLTM-24 showed 204.36% in the same trait. Total soluble solids were notably enhanced by NBLTM-4 × NBLTM-21 at 40.45%. The thickness of pericarp also revealed significant positive heterosis, with NBLTM-4 × NBLTM-28 achieving 61.29%. However, several traits experienced negative heterosis, particularly size at blossom end and number of locules, with NBLTM-3 × NBLTM-28 showing -85.60% and NBLTM-4 × NBLTM-21 at -66.67%, indicating a need for further investigation into these specific crosses. Overall, the data highlights the potential for hybridization to enhance important agricultural traits while also identifying areas requiring further research to mitigate negative effects.

Overall heterotic status of hybrids

It is important to determine the overall heterotic status of the hybrids across the characters in addition to determining the overall general combining ability status of parents and overall specific combining ability status of crosses. Hence, the method (Descending order) which was used for determining overall sca status of hybrids was also utilized to compute overall heterotic status of each cross and the results of the same are tabulated in (Table 73). From the (Table 73) it is evident that, 12 out of 25 hybrids manifested high (H) overall heterotic status while the remaining 13 crosses expressed low (L) overall heterotic status across the traits. Of these, the cross NBLTM -4 × NBLTM- 29 (H × L) manifested highest heterotic status over final norm of 245, followed by NBLTM-3 × NBLTM-28 (H × H) with 235. Among the lines, NBLTM -16 with high (H) GCA status produced maximum of four hybrids with high heterotic status. Among the testers, NBLTM-24 and NBLTM-26 with high (H) overall gca status, when crossed with all five lines produced four hybrids with high overall heterotic status. Approximately, similar type of results obtained by Nayana et al. (2021) in tomato.

It is an evident that the number of hybrids with low (L) overall heterotic status was more in H × H, L × H or H × L type of crosses than in L × L combinations. Similar type of observations was also reported by Nayana et al. (2021) in tomato. This clearly indicated the need for using the parents having high overall gca status or at least using the parent having high gca status to produce hybrids with overall heterotic status.

Summary

The study highlighted significant genetic potential in the evaluated lines and testers for breeding. Lines NBLTM-3, NBLTM-4, and NBLTM-16, along with testers NBLTM-21, NBLTM-24, NBLTM-26, and NBLTM-28, showed high General Combining Ability (GCA). Notably, 68% of the hybrids exhibited high Specific Combining Ability (SCA) across fourteen traits, indicating effective combinations and a predominance of non-additive gene action.

The analysis revealed that line contributions were greater than those of testers, with line × tester interactions also playing a crucial role. Twelve hybrids demonstrated high overall heterotic status, with NBLTM-4 × NBLTM-29 and NBLTM-3 × NBLTM-28 leading in heterotic responses. These findings suggest a promising avenue for developing superior hybrids to boost agricultural productivity.

The analysis of General Combining Ability (GCA) and Specific Combining Ability (SCA) across various traits highlights the genetic strengths of the evaluated lines and testers. Significant GCA effects show that some parental lines consistently contribute positively to trait performance, making them valuable for breeding. Notable SCA effects indicate that specific hybrid combinations can lead to exceptional results, emphasizing the importance of selecting the right parents for hybridization.

Heterosis assessments revealed both positive and negative results. Significant positive heterosis was observed in traits like fruit weight, total soluble solids, and fruit firmness, particularly in hybrids such as NBLTM-16 × NBLTM-26. This suggests that these hybrids can outperform their parents, enhancing agricultural productivity. However, negative heterosis was noted in traits like size at blossom end and number of locules, which requires careful management.

In summary, the findings underline the importance of both parental quality and hybrid selection in improving crop traits. By focusing on the strengths of lines and testers, breeders can develop superior hybrids that boost yield and quality, meeting the needs of agricultural production.

Declaration or Conflict of interest

All the authors are agreed the manuscript and there is a compatibility between the members.

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Table 1a. Performance of tomato lines for yield and yield contributing characteristics

Lines	Weight of fruits (g)	TSS (Brix)	Fruit length (cm)	Fruit width(cm)	Thickness of pericarp (mm)	Size of locules	Fruit firmness (kg/cm²)
NBLTM-3	37.85	4.45	6.25	4.95	0.775	3.85	8.13
NBLTM-4	40.80	4.15	4.30	5.95	0.725	2.45	10.86
NBLTM-16	37.80	4.45	4.55	6.55	0.605	1.65	2.73
NBLTM-17	48.95	4.15	6.45	5.15	0.675	2.15	7.23
NBLTM-23	81.35	5.10	4.15	5.45	0.575	2.35	7.56
SEm±	2.517	0.035	0.068	0.122	0.0168	0.0566	0.263
CD @ 5%	5.195	0.073	0.139	0.251	0.0347	0.1169	0.543
CD @ 1%	7.039	0.099	0.189	0.340	0.0471	0.1584	0.735

Table 1b. Performance of tomato lines for yield and yield contributing characteristics

Lines	Size at blossom end(mm)	Number of locules	Length of fruit stalk (cm)	Seed yield per fruit (mg)	Number of seeds per fruit	Germination (%)	Electrical conductivity (μS/cm/g)
NBLTM-3	0.33	2.00	3.45	130.00	76.50	77.50	54.95
NBLTM-4	0.13	4.00	1.90	228.50	43.50	75.50	68.81
NBLTM-16	0.27	5.00	2.65	323.50	44.50	37.50	48.66
NBLTM-17	0.13	3.00	2.95	165.00	34.00	64.50	31.79
NBLTM-23	0.13	3.00	2.25	293.50	44.50	79.50	51.68
SEm±	0.007	0.000	0.031	32.515	12.050	0.000	0.000
CD @ 5%	0.014	0.000	0.063	67.107	24.871	0.000	0.000
CD @ 1%	0.019	0.000	0.086	90.942	33.704	0.000	0.000

Table 1c. Performance of tomato testers for yield and yield contributing characteristics

Tester	Weight of fruits (g)	TSS (Brix)	Fruit length (cm)	Fruit width(cm)	Thickness of pericarp (mm)	Size of locules (cm)	Fruit firmness (lb/kg)
NBLTM-21	52.80	6.05	4.25	5.40	0.55	2.65	8.85
NBLTM-24 (Ty-6)	50.55	4.45	4.65	4.45	0.78	2.65	8.67
NBLTM-26 (Ty-2 and Ty-3)	81.55	5.50	4.50	5.45	0.48	2.25	5.18
NBLTM-28 (Ty-2)	100.05	6.20	6.25	5.55	0.88	2.75	7.64
NBLTM-29 (Ty-2, Ty-3 and Ty-5)	67.68	5.45	5.50	5.20	0.66	2.45	6.38
SEm±	2.517	0.035	0.068	0.122	0.0168	0.0566	0.263
CD @ 5%	5.195	0.073	0.139	0.251	0.0347	0.1169	0.543
CD @ 1%	7.039	0.099	0.189	0.340	0.0471	0.1584	0.735

Table 1d. Performance of tomato testers for yield and yield contributing characteristics

Tester	Size at blossom end (mm)	Number of locules	Length of fruit stalk (cm)	Seed yield per fruit (mg)	Number of seeds per fruit	Germination (%)	Electrical conductivity (μS/cm/g)
NBLTM-21	0.265	6.0	2.60	239.50	63.00	77.50	47.34
NBLTM-24 (Ty-6)	0.190	4.0	2.65	198.50	42.50	47.50	44.31
NBLTM-26 (Ty-2 and Ty-3)	0.215	4.0	2.55	220.00	33.50	75.50	48.72
NBLTM-28 (Ty-2)	0.625	2.0	2.90	144.00	46.00	94.50	33.60
NBLTM-29 (Ty-2, Ty-3 and Ty-5)	0.245	3.0	2.40	194.50	42.50	78.50	69.38
SEm±	0.007	0.000	0.031	32.515	12.050	0.000	0.000
CD @ 5%	0.014	0.000	0.063	67.107	24.871	0.000	0.000
CD @ 1%	0.019	0.000	0.086	90.942	33.704	0.000	0.000

Table 1e. Performance of tomato parental cross (hybrids) for yield and yield contributing characteristics

	Weight of fruits (g)	Total Soluble solids (Brix)	Fruit length (cm)	Fruit width (cm)	Thickness of pericarp (cm)	Size of locules (cm)	Fruit firmness (lb/kg)	Size at blossom end (mm)
NBLTM-3×NBLTM-21	96.80	3.95	4.55	4.15	0.78	3.45	11.74	0.13
NBLTM-3×NBLTM-24	75.80	4.5	6.15	4.10	0.80	2.65	11.80	0.19
NBLTM-3×NBLTM-26	67.30	5.25	5.22	4.55	0.72	3.45	6.08	0.16
NBLTM-3×NBLTM-28	64.95	4.75	5.74	3.55	0.63	2.45	5.31	0.09
NBLTM-3×NBLTM-29	53.90	4.25	5.09	4.15	0.47	1.90	6.10	0.19
NBLTM-4×NBLTM-21	97.75	6.25	4.25	5.86	0.88	3.15	6.92	0.09
NBLTM-4×NBLTM-24	74.55	6.00	5.15	4.62	1.15	3.55	7.03	0.09
NBLTM-4×NBLTM-26	67.85	4.55	4.75	3.59	1.15	4.05	8.78	0.11
NBLTM-4×NBLTM-28	106.95	3.95	4.88	6.32	1.25	3.95	10.24	0.14
NBLTM-4×NBLTM-29	44.05	3.35	3.52	4.22	0.85	1.30	5.63	0.11
NBLTM-16×NBLTM-21	69.50	5.15	3.60	5.45	0.58	2.15	9.05	0.11
NBLTM-16×NBLTM-24	115.20	4.45	5.83	6.11	0.72	1.90	9.03	0.16
NBLTM-16×NBLTM-26	138.50	5.25	6.43	7.89	0.58	1.90	8.08	0.10
NBLTM-16×NBLTM-28	116.20	5.55	6.62	7.74	0.56	2.25	9.89	0.19
NBLTM-16×NBLTM-29	111.65	5.45	5.59	6.63	0.59	2.05	6.11	0.15
NBLTM-17×NBLTM-21	48.80	4.30	5.53	4.59	0.78	3.45	10.74	0.11
NBLTM-17×NBLTM-24	45.90	3.55	4.35	3.75	0.36	1.95	9.24	0.12
NBLTM-17×NBLTM-26	80.60	4.10	5.30	4.27	0.41	2.35	10.89	0.11
NBLTM-17×NBLTM-28	45.50	4.20	5.19	4.39	0.37	2.35	7.10	0.12
NBLTM-17×NBLTM-29	43.00	4.20	3.45	2.30	0.47	1.15	9.08	0.11
NBLTM-23×NBLTM-21	51.20	4.95	3.32	3.10	0.43	1.25	6.34	0.16
NBLTM-23×NBLTM-24	40.70	4.05	3.25	3.40	0.41	2.05	5.69	0.16
NBLTM-23×NBLTM-26	45.55	4.10	3.31	3.55	0.39	2.15	8.32	0.18
NBLTM-23×NBLTM-28	38.40	3.95	2.82	3.30	0.36	1.80	5.97	0.16
NBLTM-23×NBLTM-29	72.90	4.90	4.15	4.55	0.41	1.95	7.32	0.18

Table 1f. Performance of tomato parental crosses (hybrids) for yield and yield contributing characteristics of tomato

	Number of locules	Length of fruit stalk (cm)	Seed yield per fruit (g)	Number of seeds per fruit	Seed germination (%)	Electrical conductivity (μS/cm/g)
NBLTM-3×NBLTM-21	3.00	3.20	330	92.00	81.50	33.17
NBLTM-3×NBLTM-24	3.00	2.45	235	97.00	85.50	26.49
NBLTM-3×NBLTM-26	2.00	2.75	195	44.50	84.50	28.76
NBLTM-3×NBLTM-28	2.00	3.15	85	70.00	80.50	63.58
NBLTM-3×NBLTM-29	2.00	2.80	100	24.50	87.50	48.36
NBLTM-4×NBLTM-21	2.00	2.45	355	107.00	84.50	25.65
NBLTM-4×NBLTM-24	3.00	2.45	320	100.50	75.50	40.17
NBLTM-4×NBLTM-26	3.00	2.40	140	89.00	76.50	34.51
NBLTM-4×NBLTM-28	3.00	2.75	390	124.50	87.50	23.48
NBLTM-4×NBLTM-29	3.00	2.25	140	24.00	82.50	21.27
NBLTM-16×NBLTM-21	4.00	2.45	275	114.50	72.50	19.83
NBLTM-16×NBLTM-24	2.00	2.65	290	125.00	71.50	19.72
NBLTM-16×NBLTM-26	6.00	2.95	305	166.50	70.50	22.26
NBLTM-16×NBLTM-28	5.00	2.85	290	86.00	83.50	21.72
NBLTM-16×NBLTM-29	4.00	2.85	210	63.50	74.50	25.22
NBLTM-17×NBLTM-21	3.00	2.65	135	59.00	88.50	18.42
NBLTM-17×NBLTM-24	4.00	2.85	146	63.00	87.50	24.10
NBLTM-17×NBLTM-26	3.00	2.75	110	42.50	84.50	29.90
NBLTM-17×NBLTM-28	2.00	2.95	120	38.00	86.50	23.82
NBLTM-17×NBLTM-29	2.00	2.75	80	31.50	87.50	26.85
NBLTM-23×NBLTM-21	4.00	2.55	295	115.50	73.50	25.67
NBLTM-23×NBLTM-24	3.00	2.65	150	49.00	76.50	27.41
NBLTM-23×NBLTM-26	3.00	2.65	115	64.00	70.50	22.67
NBLTM-23×NBLTM-28	3.00	2.45	215	83.00	77.50	24.11
NBLTM-23×NBLTM-29	5.00	2.55	195	63.50	76.50	27.01

Table 2a. ANOVA for combining ability

Source of variation	DF	Weight of fruits	TSS	Fruit length	Fruit width	Thickness of pericarp	Size of locules	Fruit firmness	Size at blossom end
Replications	1	18.422	0.017	0.006	0.001	0.000	0.129**	0.042	0.000
Treatments	34	1467.23**	1.161**	2.169**	3.233**	0.104**	1.135**	8.583**	0.019**
Parents	9	932.99**	1.191**	1.723**	0.635**	0.030**	0.636**	9.786**	0.044**
Parents vs. Crosses	1	2268.90**	2.413**	1.884**	8.352**	0.009	0.132*	17.155**	0.187**
Crosses	24	1634.16**	1.098**	2.347**	3.993**	0.136**	1.365**	8.130**	0.002**
Lines	4	5906.65**	1.785	7.668**	16.556**	0.648**	3.1	9.667	0.005*
Testers	4	295.99	0.431	1.472	0.816	0.028	1.961	6.957	0.001
Lines × Testers	16	900.584**	1.093**	1.236**	1.647**	0.036**	0.782**	8.038**	0.002*
Error	34	53.513	0.0.13	0.106	0.06	0.001	0.014	0.465	0.00

Table 2b. ANOVA for combining ability

Source of variation	DF	Number of locules	Length of fruit stalk	Seed yield/fruit	Number of seeds/fruit	Germination	Electrical conductivity
Replications	1	0.000	0.302**	30.229	248.914	17.5**	0.700**
Treatments	34	2.538	0.172**	14322.92**	2308.65**	238.867	420.461**
Parents	9	3.200**	0.352**	7596.37	343.05	560.467**	311.137**
Parents vs. Crosses	1	2.766**	0.048*	337.423	13245.75**	1289.286**	6763.008**
Crosses	24	2.28	0.110**	17428.11**	2590.042**	74.5	197.184**
Lines	4	5.28	0.283	41736.28**	5878.9	300.800**	510.06
Testers	4	0.28	0.074	26606.28	4544.65	49.6	65.761
Lines × Testers	16	2.03	0.076**	9056.53	1279.18	24.15	151.821**
Error	34	0.00	0.005	3785.2	550.356	0.000	0.000

Table 3a. Estimation of gca effects on lines of tomato

Lines	Weight of fruits (g)	TSS (Brix)	Fruit length (cm)	Fruit width (cm)	Thickness of pericarp (mm)	Size of locules (cm)	Fruit firmness (lb/kg)
NBLTM-3	-0.790	-0.114**	0.629**	-0.545**	0.036**	0.356**	0.106
NBLTM-4	5.690*	0.236**	-0.212**	0.276*	0.411**	0.776**	-0.380
NBLTM-16	37.670**	0.586**	0.891**	2.121**	-0.037*	-0.374**	0.332
NBLTM-17	-19.780**	-0.514**	0.044	-0.785**	-0.166**	-0.174**	1.312**
NBLTM-23	-22.790**	-0.194**	-1.353**	-1.065**	-0.245**	-0.584**	-1.371**
SEm±	1.7797	0.025	0.048	0.086	0.012	0.040	0.186
CD @ 5%	3.673	0.052	0.098	0.178	0.025	0.083	0.384
CD @ 1%	4.978	0.07	0.133	0.241	0.033	0.112	0.520

Table 3 b. Estimation of gca effects on linesof tomato

Lines	Size at blossom end (mm)	Number of locules	Length of fruit stalk (cm)	Seed yield per fruit (mg)	Number of seeds per fruit	Germination (%)	Electrical conductivity (μS/cm/g)
NBLTM-3	0.014**	-0.760**	0.182**	-19.840	-11.90	3.60**	11.906**
NBLTM-4	-0.027**	-0.360**	-0.228**	60.160	11.50	1.00**	0.850**
NBLTM-16	0.003	1.040**	0.062*	65.160*	33.60**	-5.80**	-6.416**
NBLTM-17	-0.020**	-0.360**	0.102**	-90.640*	-30.70	6.60**	-3.548**
NBLTM-23	0.028**	0.440**	-0.118**	-14.840	-2.50	-5.40**	-2.792**
SEm±	0.0049	0.000	0.0217	22.99	8.521	0.000	0.000
CD @ 5%	0.0101	0.000	0.048	47.45	17.586	0.000	0.000
CD @ 1%	0.0137	0.000	0.067	64.306	23.83	0.000	0.000

Table 3 c. Estimation of gca effects on tester of tomato

Tester	Weight of fruits (g)	TSS (Brix)	Fruit length (cm)	Fruit width (cm)	Thickness of pericarp (mm)	Size of locules (cm)	Fruit firmness (lb/kg)
NBLTM-21	0.270	0.336**	-0.471**	-0.015	0.043*	0.266**	0.858**
NBLTM-24	-2.110	-0.144**	0.224**	-0.250*	0.045**	-0.004	0.459
NBLTM-26	7.420**	0.066	0.281**	0.125	0.006	0.356**	0.332
NBLTM-28	1.860	-0.104**	0.327**	0.416**	-0.011	0.136*	-0.398
NBLTM-29	-7.440**	-0.154**	-0.362**	-0.274*	-0.084**	-0.754**	-1.252**
SEm±	1.7797	0.025	0.048	0.086	0.012	0.040	0.186
CD @ 5%	3.673	0.052	0.098	0.178	0.025	0.083	0.384
CD @ 1%	4.978	0.07	0.133	0.241	0.033	0.112	0.520

Table 3d. Estimation of gca effects on tester of tomato

Tester	Size at blossom end (mm)	Number of locules	Length of fruit stalk (cm)	Seed yield per fruit (mg)	Number of seeds per fruit	Germination (%)	Electrical conductivity (μS/cm/g)
NBLTM-21	-0.017*	0.040**	-0.028	69.160*	20.100	-0.200**	-3.618**
NBLTM-24	0.004	-0.160**	-0.078*	19.360	9.400	-1.000**	-0.588**
NBLTM-26	-0.005	0.240**	0.012	-35.840	3.800	-3.000**	-0.546**
NBLTM-28	0.005	-0.160**	0.142**	11.160	2.800	2.800**	3.176**
NBLTM-29	0.011	0.040**	-0.048	-63.840	-36.100**	1.400**	1.576**
SEm±	0.0049	0.000	0.0217	22.99	8.521	0.000	0.000
CD @ 5%	0.0101	0.000	0.048	47.45	17.586	0.000	0.000
CD @ 1%	0.0137	0.000	0.067	64.306	23.83	0.000	0.000

Table 4a. Overall general combining ability status of testers of tomato

Testers	Total rank	Overall GCA status
NBLTM-21	41	H
NBLTM-24	45	H
NBLTM-26	37	H
NBLTM-28	42	H
NBLTM-29	60	L

Final norm: 45

Table 4b. Overall general combining ability status of lines of tomato

Lines	Total rank	Overall GCA status
NBLTM-3	39	H
NBLTM-4	42	H
NBLTM-16	30	H
NBLTM-17	52	L
NBLTM-23	61	L

Final norm: 44.8

H: High overall general combiner, L: Low overall general combiner

Note: Arranged data in ascending order

Table 5 a. Estimates of specific combining ability effects (sca) in crosses for yield and yield contributing characteristics of tomato

Hybrids	Weight of fruits (g)	TSS (Brix)	Fruit length (cm)	Fruit width (cm)	Thickness of pericarp (mm)	Size of locules (cm)	Fruit firmness (lb/kg)	Size at blossom end (mm)
NBLTM-3×NBLTM-21	24.780**	-0.856**	-0.330*	0.065	0.052	0.404**	2.672**	-0.009
NBLTM-3×NBLTM-24	6.160	-0.176*	0.575**	0.250	0.075*	-0.126	3.136**	0.035*
NBLTM-3×NBLTM-26	-11.870**	0.714**	-0.407**	0.325	0.039	0.314*	-2.457**	0.014
NBLTM-3×NBLTM-28	-8.660	0.384**	0.067	-0.966**	-0.044	-0.466**	-2.497**	-0.066**
NBLTM-3×NBLTM-29	-10.410	-0.066	0.096	0.324	-0.121**	-0.126	-0.853	0.028
NBLTM-4×NBLTM-21	19.250**	1.094**	0.211	0.954**	-0.223**	-0.316*	-1.657**	-0.003
NBLTM-4×NBLTM-24	-1.570	1.324**	0.416**	-0.056	0.050	0.354**	-1.153	-0.024
NBLTM-4×NBLTM-26	-17.800**	-0.336**	-0.041	-1.456**	0.089	0.494**	0.724	0.010
NBLTM-4×NBLTM-28	26.860**	-0.766**	0.038	0.978**	0.206**	0.614**	2.919**	0.025
NBLTM-4×NBLTM-29	-26.740**	-1.316**	-0.623	-0.422	-0.121**	-1.146**	-0.832	-0.006
NBLTM-16×NBLTM-21	-40.980**	-0.356**	-1.542**	-1.301**	-0.075*	-0.166	-0.239	-0.008
NBLTM-16×NBLTM-24	7.100	-0.576**	-0.007	-0.406	0.073	-0.146	0.145	0.011
NBLTM-16×NBLTM-26	20.870**	0.014	0.536**	1.004**	-0.038	-0.506**	-0.683	-0.040*
NBLTM-16×NBLTM-28	4.130	0.484**	0.675**	0.563*	-0.031	0.064	1.852**	0.045**
NBLTM-16×NBLTM-29	8.880	0.434**	0.339*	0.138	0.072	0.754**	-1.074	-0.006
NBLTM-17×NBLTM-21	-4.230	-0.106	1.240**	0.745**	0.254**	0.934**	0.476	0.015
NBLTM-17×NBLTM-24	-4.750	-0.376**	-0.640**	0.140	-0.158**	-0.296*	-0.630	-0.006
NBLTM-17×NBLTM-26	20.420**	-0.036	0.253	0.280	-0.079*	-0.256*	1.152	0.003
NBLTM-17×NBLTM-28	-9.120	0.234**	0.102	0.119	-0.097*	-0.036	-1.918**	0.003
NBLTM-17×NBLTM-29	-2.320	0.284**	-0.954**	-1.286**	0.081*	-0.346**	0.921	-0.013
NBLTM-23×NBLTM-21	1.180	0.224**	0.422**	-0.465	-0.007	-0.856**	-1.251*	0.007
NBLTM-23×NBLTM-24	-6.940	-0.196*	-0.343*	0.070	-0.039	0.214	-1.497*	-0.014
NBLTM-23×NBLTM-26	-11.620*	-0.356**	-0.340*	-0.155	-0.010	-0.046	1.265*	0.015
NBLTM-23×NBLTM-28	-13.210*	-0.336**	-0.881**	-0.696*	-0.033	-0.176	-0.355	-0.005
NBLTM-23×NBLTM-29	30.590**	0.664**	1.143**	1.244**	0.090*	0.864**	1.839**	-0.001
SE±m	3.9794	0.0579	0.1067	0.1923	0.0266	0.0895	0.4155	0.0109
CD (P=0.05)	8.2132	0.1154	0.2201	0.3968	0.0549	0.1848	0.8576	0.0225
CD (P=0.01)	11.1303	0.1564	0.2983	0.5378	0.0745	0.2504	1.1622	0.0306

Table 5 b. Estimates of specific combining ability effects (sca) in crosses for yield and yield contributing characteristics of tomato

Hybrids	Number of locules	Length of fruit stalk (cm)	Seed yield / fruit (mg)	Number of seeds /fruit	Germination (%)	Electrical conductivity (μS/cm/g)
NBLTM-3×NBLTM-21	0.560**	0.358**	71.840	6.300	-2.200**	-3.284**
NBLTM-3×NBLTM-24	0.760**	-0.342**	26.640	22.000	2.600**	-12.994**
NBLTM-3×NBLTM-26	-0.640**	-0.132	41.840	-24.900	3.600**	-10.766**
NBLTM-3×NBLTM-28	-0.240**	0.138*	-115.160	1.600	-6.200**	20.322**
NBLTM-3×NBLTM-29	-0.440**	-0.022	-25.160	-5.000	2.200**	6.712**
NBLTM-4×NBLTM-21	-0.840**	0.018	16.840	-2.100	3.400**	0.252**
NBLTM-4×NBLTM-24	0.360**	0.068	31.640	2.100	-4.800**	11.742**
NBLTM-4×NBLTM-26	-0.040**	-0.072	-93.160	-3.800	-1.800**	6.040**
NBLTM-4×NBLTM-28	0.360**	0.148*	109.840	32.700	3.400**	-8.712**
NBLTM-4×NBLTM-29	0.160**	-0.162*	-65.160	-28.900	-0.200**	-9.322**
NBLTM-16×NBLTM-21	-0.240**	-0.272**	-68.160	-16.700	-1.800**	1.698**
NBLTM-16×NBLTM-24	-2.040**	-0.022	-3.360	4.500	-2.000**	-1.442**
NBLTM-16×NBLTM-26	1.560**	0.188**	66.840	51.600	-1.000**	1.056**
NBLTM-16×NBLTM-28	0.960**	-0.042	4.840	-27.900	6.200**	-3.206**
NBLTM-16×NBLTM-29	-0.240**	0.148*	-0.160	-11.500	-1.400**	1.894**
NBLTM-17×NBLTM-21	0.160**	-0.112	-52.360	-7.900	1.800**	-2.580**
NBLTM-17×NBLTM-24	1.360**	0.138*	8.440	6.800	1.600**	0.070**
NBLTM-17×NBLTM-26	-0.040**	-0.052	27.640	-8.100	0.600**	5.828**
NBLTM-17×NBLTM-28	-0.640**	0.018	-9.360	-11.600	-3.200**	-3.974**
NBLTM-17×NBLTM-29	-0.840**	0.008	25.640	20.800	-0.800**	0.656**
NBLTM-23×NBLTM-21	0.360**	0.008	31.840	20.400	-1.200**	3.914**
NBLTM-23×NBLTM-24	-0.440**	0.158*	-63.360	-35.400	2.600**	2.624**
NBLTM-23×NBLTM-26	-0.840**	0.068	-43.160	-14.800	-1.400**	-2.158**
NBLTM-23×NBLTM-28	-0.440**	-0.262**	9.840	5.200	-0.200**	-4.440**
NBLTM-23×NBLTM-29	1.360**	0.028	64.840	24.600	0.200**	0.060**
SEm±	0.000	0.0486	51.410	19.0531	0.000	0.000
CD (P=0.05)	0.000	0.1002	106.106	39.3240	0.000	0.000
CD (P=0.01)	0.000	0.1358	143.792	53.2908	0.000	0.000

Table 6. Overall specific combining ability status of lines and testers of tomato

Tester	Lines
Tester	NBLTM-3 (H)	NBLTM-4 (H)	NBLTM-16 (H)	NBLTM-17 (L)	NBLTM-23 (L)
NBLTM-21 (H)	209 (L)	200 (L)	168 (H)	169 (H)	166 (H)
NBLTM-24 (H)	206 (L)	163 (H)	141 (H)	164 (H)	121 (H)
NBLTM-26 (H)	177 (L)	161 (H)	202 (L)	152 (H)	142 (H)
NBLTM-28 (H)	162 (H)	268 (L)	185 (L)	155 (H)	164 (H)
NBLTM-29 (L)	135 (H)	161 (H)	146 (H)	170 (H)	226 (L)

Final norm : 172.52 H: High overall general combiner

L: Low overall general combiner

Note : Arranged the data in Ascending order

Table 7. Estimates of variance components for different traits of tomato

Characteristics	σ²GCA	σ²SCA	σ²GCA/ σ²SCA
Weight of the fruits	27.509	434.456	0.0633
Total soluble solids	0.0002	0.5435	0.0004
Fruit length	0.0417	0.6067	0.0687
Fruit width	0.0880	0.7886	0.1116
Thickness of pericarp	0.0038	0.0171	0.2222
Size of locules	0.0219	0.3828	0.0572
Fruit firmness	0.0034	3.8464	0.0009
Size at blossom end	0.0000	0.0007	0.0000
Number of locules	0.0094	1.0150	0.0093
Length of fruit stalk	0.0013	0.0356	0.0365
Seed yield/fruit	313.934	1885.275	0.1665
Number of seeds/fruit	49.1575	276.5667	0.1777
Germination	1.8881	12.0750	0.1564
Electrical conductivity	1.7011	75.9104	0.0224

Table 8. Proportional contribution of lines, testers and line × tester interaction towards total variance(in percentage) in tomato

Characteristics	Line	Tester	Line × Tester
Weight of the fruits	60.24	3.02	36.74
Total soluble solids	27.09	6.55	66.37
Fruit length	54.44	10.45	35.11
Fruit width	69.10	3.41	27.50
Thickness of pericarp	79.21	3.39	17.41
Size of locules	37.87	23.95	38.18
Fruit firmness	19.82	14.26	65.92
Size at blossom end	41.13	9.23	49.64
Number of locules	38.60	2.05	59.36
Length of fruit stalk	42.86	11.20	45.98
Seed yield/fruit	39.91	25.44	34.64
Number of seeds/fruit	37.83	29.24	32.93
Germination	67.29	11.10	21.61
Electrical conductivity	43.11	5.56	51.33

Table 9 a. Estimates of mid-parent, better parent and standard check heterosis of 25 crosses for weight of fruit, total soluble solids and fruit length of tomato

SCH	Weight of fruits			Total Soluble solids			Fruit length
SCH	Mid	Better	Standard	Mid	Better	Standard	Mid	Better	Standard
NBLTM-3×NBLTM-21	113.57*	83.33*	115.75*	-24.76*	-34.71*	-11.24*	-13.33*	-27.20*	-27.20*
NBLTM-3×NBLTM-24	71.49*	49.95*	100.26*	-6.74*	-6.74*	-6.74*	12.84*	-1.60	-1.60
NBLTM-3×NBLTM-26	12.73	-17.47	77.81*	5.53*	-4.55*	17.98*	-2.79	-16.40*	-16.40*
NBLTM-3×NBLTM-28	-5.80	-35.08*	71.6*	-10.80*	-23.39*	6.74*	-8.08*	-8.08*	-8.08*
NBLTM-3×NBLTM-29	2.16	-20.35	42.4*	-14.14*	-22.02*	-4.49	-13.45*	-18.64*	-18.64*
NBLTM-4×NBLTM-21	108.87*	85.13*	158.26*	22.55*	3.31	40.45*	-0.58	-1.16	-32.00
NBLTM-4×NBLTM-24	63.22*	47.48*	96.96*	39.53*	34.83*	34.83*	15.08*	10.75*	-17.60*
NBLTM-4×NBLTM-26	10.91	-16.80	79.26*	-5.70*	-17.27*	2.25	7.95	5.56	-24.00*
NBLTM-4×NBLTM-28	51.86*	6.90	182.56*	-23.67*	-36.29*	-11.24*	-7.58*	-22.00*	-22.00*
NBLTM-4×NBLTM-29	-18.78	-34.91*	16.38	-30.21*	-38.53*	-24.72*	-28.06*	-35.91*	-43.40*
NBLTM-16×NBLTM-21	53.42*	31.63*	83.62*	-1.90	-14.88*	15.73*	-18.18*	-20.88*	-42.40*
NBLTM-16×NBLTM-24	160.78*	127.89*	204.36*	0.00	0.00	0.00	26.74*	25.38*	-6.72
NBLTM-16×NBLTM-26	132.09*	69.83*	265.92*	5.53*	-4.55*	17.98*	42.10*	41.32*	2.88
NBLTM-16×NBLTM-28	68.59*	16.14*	207*	4.23*	-10.48*	24.72*	22.50*	5.84	5.84
NBLTM-16×NBLTM-29	111.71*	64.98*	194.98*	10.10*	0.00	22.47*	11.24*	1.64	-10.56*
NBLTM-17×NBLTM-21	-4.08	-7.58	28.93	-15.69*	-28.93*	-3.37	3.46	-14.19*	-11.44*
NBLTM-17×NBLTM-24	-7.74	-9.20	21.27	-17.44*	-20.22*	-20.22*	-21.62*	-32.56*	-30.40*
NBLTM-17×NBLTM-26	23.52*	-1.16	112.95*	-15.03*	-25.45*	-7.87*	-3.20	-17.83*	-15.20*
NBLTM-17×NBLTM-28	-38.93*	-54.52*	20.21	-18.84*	-32.26*	-5.62*	-18.19*	-19.46*	-16.88*
NBLTM-17×NBLTM-29	-26.26*	-36.46*	13.61	-12.50*	-22.94*	-5.62*	-42.26*	-46.51*	-44.80*
NBLTM-23×NBLTM-21	-23.67*	-37.06*	35.27	-11.21*	-18.18*	11.24*	-20.95*	-21.88*	-46.88*
NBLTM-23×NBLTM-24	-38.29*	-49.97*	7.53	-15.18*	-20.59*	-8.99*	-26.14*	-30.11*	-48.00*
NBLTM-23×NBLTM-26	-44.08*	44.14*	20.34	-22.64*	-25.45*	-7.87*	-23.47*	-26.44*	-47.04*
NBLTM-23×NBLTM-28	-57.66*	61.62*	1.45	-30.09*	-36.29*	-11.24*	-45.87	-54.96*	-54.96*
NBLTM-23×NBLTM-29	-2.16	10.39	92.60*	-7.11*	-10.09*	10.11*	-13.99*	-24.55*	-33.60*

Table 9 b. Estimates of mid-parent, better parent and standard check heterosis of 25 crosses for fruit width, thickness of pericarp and size of locules in tomato

SCH	Fruit width			Thickness of pericarp			Size of locules
SCH	Mid	Better	Standard	Mid	Better	Standard	Mid	Better	Standard
NBLTM-3×NBLTM-21	-19.81*	-23.15*	-16.16*	17.42*	0.00	0.00	6.15	-10.39*	-10.39*
NBLTM-3×NBLTM-24	-12.77	-17.17*	-17.17*	3.23	3.23	3.23	-18.46*	31.17*	-31.17*
NBLTM-3×NBLTM-26	-12.50	-16.51*	-8.08	16.00*	-6.45	-6.45	13.11*	-10.39*	-10.39*
NBLTM-3×NBLTM-28	-32.38*	-36.04*	-28.28*	-24.24*	-28.57*	-19.35*	-25.76*	-36.36*	-36.36*
NBLTM-3×NBLTM-29	-18.23*	-20.19*	-16.16*	-33.80*	-38.71*	-38.71*	-39.68*	-50.65*	-50.65*
NBLTM-4×NBLTM-21	3.26	-1.51	18.38*	37.80*	20.69*	12.90	23.53*	18.87*	-18.18*
NBLTM-4×NBLTM-24	-11.25	-22.44*	-6.77	53.33*	48.39*	48.39*	39.22*	33.96*	-7.79
NBLTM-4×NBLTM-26	-37.02*	-39.66*	-27.47*	91.67*	58.62*	48.39*	72.34*	65.31*	5.19
NBLTM-4×NBLTM-28	9.83	6.13	27.58*	56.25*	42.86*	61.29*	51.92*	43.64*	2.60
NBLTM-4×NBLTM-29	-24.22*	-28.99*	-14.65	22.74*	17.24*	9.68	-46.94*	-46.94*	-66.23*
NBLTM-16×NBLTM-21	-8.79	-16.79*	10.10	0.00	-4.96	-25.81*	0.00	-18.87*	-44.16*
NBLTM-16×NBLTM-24	11.09	-6.72	23.43*	5.07	-6.45	-6.45	-11.63	-28.30*	-50.65*
NBLTM-16×NBLTM-26	31.58*	20.53*	59.49*	6.48	-4.96	-25.81*	-2.56	-15.56	-50.65*
NBLTM-16×NBLTM-28	28.02*	18.24*	56.46*	-23.65*	-35.43*	-27.10*	2.27	-18.18*	-41.56*
NBLTM-16×NBLTM-29	12.85*	1.22	33.94*	-5.93	-9.85	-23.23*	0.00	-16.33*	-46.75*
NBLTM-17×NBLTM-21	-12.99*	-15.00*	-7.27	27.05*	14.81	0.00	43.75*	30.19*	-10.39*
NBLTM-17×NBLTM-24	-21.87*	-27.18*	-24.24*	-49.66*	-52.90*	-52.90*	-18.75*	-26.42*	-49.35*
NBLTM-17×NBLTM-26	-19.53*	-21.74*	-13.84	-29.57*	-40.00	-47.74*	6.82	4.44	-38.96*
NBLTM-17×NBLTM-28	-17.85*	-20.81*	-11.21	-52.26*	-57.71*	-52.26*	-4.08	-14.55*	-38.96*
NBLTM-17×NBLTM-29	-55.56	-55.77*	-53.54*	-28.84*	-29.63*	-38.71*	-50.00*	-53.06*	-70.13*
NBLTM-23×NBLTM-21	-42.86*	-43.12*	-37.37*	-22.32*	-24.35*	-43.87*	-50.00*	-52.83*	-67.53*
NBLTM-23×NBLTM-24	-31.31	-37.61*	-31.31	-40.00*	-47.74*	-47.74*	-18.00*	-22.64	-46.75*
NBLTM-23×NBLTM-26	-34.86*	-34.86*	-28.28*	-24.76*	-31.30*	-49.03*	-6.52	-8.51	-44.16*
NBLTM-23×NBLTM-28	-40.00*	-40.54*	-33.33*	-51.03*	-59.43*	-54.19*	-29.41*	-34.55*	-53.25*
NBLTM-23×NBLTM-29	-14.55	-16.51*	-8.08	-34.41*	-38.64*	-47.74*	-18.75	-20.41*	-49.35*

Table 9 c. Estimates of mid-parent, better parent and standard check heterosis of 25 crosses for fruit firmness, size at blossom end and number of locules in tomato

SCH	Fruit firmness			Size at blossom end			Number of locules
SCH	Mid	Better	Standard	Mid	Better	Standard	Mid	Better	Standard
NBLTM-3×NBLTM-21	38.26*	32.60*	44.43*	-57.60*	-61.54*	-61.54*	-25.00*	-50.00*	50.00*
NBLTM-3×NBLTM-24	40.52*	36.10*	45.23*	-26.21*	-41.54*	-41.54*	0.00	-25.00*	50.00*
NBLTM-3×NBLTM-26	-8.61	-25.17*	-25.17*	-40.74*	-50.77*	-50.77*	-33.33*	-50.00*	0.00
NBLTM-3×NBLTM-28	-32.61*	-34.65*	-34.65*	-80.05*	-85.60*	-72.31*	0.00	0.00	0.00
NBLTM-3×NBLTM-29	-15.89	-24.92*	-24.92*	-33.33*	-41.54*	-41.54*	-20.00*	-33.33*	0.00
NBLTM-4×NBLTM-21	-29.78*	-36.28*	-14.83	-53.85*	-66.04*	-72.31*	-60.00*	-66.67*	0.00
NBLTM-4×NBLTM-24	-28.06*	-35.31*	-13-54	-42.86*	-52.63*	-72.31*	-25.00*	-25.00*	50.00*
NBLTM-4×NBLTM-26	9.41	-19.20*	8.00	-32.35*	-46.51*	-64.62*	-25.00*	-25.00*	50.00*
NBLTM-4×NBLTM-28	10.73	-5.71	26.03*	-62.67*	-77.60*	-56.92*	0.00	-25.00*	50.00*
NBLTM-4×NBLTM-29	-34.63*	-48.11*	-30.65*	-37.84*	-53.06*	-64.62*	-14.29*	-25.00*	50.00*
NBLTM-16×NBLTM-21	56.30*	2.26	11.38	-57.01*	-57.41*	-64.62*	-27.27*	-33.33*	100.00*
NBLTM-16×NBLTM-24	58.51*	4.21	11.20	-32.61*	-42.59*	-52.31*	-55.56*	-60.00*	0.00
NBLTM-16×NBLTM-26	104.30*	55.98*	-0.55	-60.82*	-64.81*	-70.77*	33.33*	20.00*	200.00*
NBLTM-16×NBLTM-28	90.74*	29.47*	21.66*	-57.54*	-69.60*	-41.54*	42.86*	0.00	150.00*
NBLTM-16×NBLTM-29	34.03*	-4.31	-24.86*	-43.69*	-46.30*	-55.38*	0.00	-20.00*	100.00*
NBLTM-17×NBLTM-21	33.64*	21.41*	32.25*	-1.03*	-56.60*	-64.62*	-33.33*	-50.00*	50.00*
NBLTM-17×NBLTM-24	16.23	6.57	13.72	-26.98*	-39.47*	-64.62*	14.29*	0.00	100.00*
NBLTM-17×NBLTM-26	75.58*	50.69*	34.09*	-32.35*	-46.51*	-64.62*	-14.29*	-25.00*	50.00*
NBLTM-17×NBLTM-28	-4.54	-7.07	-12.68	-66.67*	-80.00*	-61.54*	-20.00*	-33.33*	0.00
NBLTM-17×NBLTM-29	33.43*	25.59*	11.75	-37.84*	-53.06*	-64.62*	-33.33*	-33.33*	0.00
NBLTM-23×NBLTM-21	-22.77*	-28.42*	-22.03*	-20.51*	-41.51*	-52.31*	-11.11*	-33.33*	100.00*
NBLTM-23×NBLTM-24	-29.86*	-34.37*	-29.97*	-1.59	-18.42	-52.31*	-14.29*	-25.00*	50.00*
NBLTM-23×NBLTM-26	30.74*	10.19	2.46	2.94	-18.60	-46.15*	-14.29*	-25.00*	50.00*
NBLTM-23×NBLTM-28	-21.33*	-21.74*	-26.46*	-56.00	-73.60*	-49.23*	20.00*	0.00	50.00*
NBLTM-23×NBLTM-29	4.99	-3.18	-9.97	-5.41	-28.57*	-46.15*	66.67*	66.67*	150.00*

Table 9 d. Estimates of mid-parent, better parent and standard check heterosis of 25 crosses for length of fruit stalk, seed yield per fruitand number of seeds per fruit in tomato

SCH	Length of fruit stalk			Seed yield per fruit			Number of seeds per fruit
SCH	Mid	Better	Standard	Mid	Better	Standard	Mid	Better	Standard
NBLTM-3×NBLTM-21	5.79*	-7.25*	-7.25*	78.62	37.79	153.85	31.90	20.26	20.26
NBLTM-3×NBLTM-24	-19.67*	-28.99*	-28.99*	43.07	18.39	80.77	63.03	26.80	26.80
NBLTM-3×NBLTM-26	-8.33*	-20.29*	-20.29*	11.43	-11.36	50.00	-19.09	-41.83	-41.83
NBLTM-3×NBLTM-28	-0.79	-8.70*	-8.70*	-37.96	-40.97	-34.62	14.29	-8.50	-8.50
NBLTM-3×NBLTM-29	-4.27	-18.84*	-18.84*	-38.37	-48.59	-23.08	-58.82	-67.97	67.97
NBLTM-4×NBLTM-21	8.89*	-5.77	-28.99*	51.71	48.23	173.08*	100.94	69.84	39.87
NBLTM-4×NBLTM-24	7.69*	-7.55*	-28.99*	49.88	40.04	146.15	133.72	131.03.	31.37
NBLTM-4×NBLTM-26	7.87*	-5.88	-30.43*	-37.57	-38.73	7.69	131.17	104.60	16.34
NBLTM-4×NBLTM-28	14.58*	-5.17	-20.29*	109.40*	70.68	200.00	178.21*	170.65*	62.75
NBLTM-4×NBLTM-29	4.65	-6.25	-34.78*	-33.81	-38.73	7.69	-44.19	-44.83	-68.63
NBLTM-16×NBLTM-21	-6.67*	-7.55*	-28.99*	-2.31	-14.99	111.54	113.02	81.75	49.67
NBLTM-16×NBLTM-24	0.00	0.00	-23.19*	11.11	-10.36	123.08	187.36*	180.90*	63.40
NBLTM-16×NBLTM-26	13.46*	11.32*	-14.49*	12.24	-5.72	134.62	162.92*	274.16*	117.65*
NBLTM-16×NBLTM-28	2.70	-1.72	-17.39*	24.06	-10.36	123.08	90.06	86.96	12.42
NBLTM-16×NBLTM-29	12.87*	7.55*	-17.39*	-18.92	-35.09	61.54	45.98	42.70	-16.99
NBLTM-17×NBLTM-21	-4.50	-10.17*	-23.19*	-33.25	-43.63	3.85	21.65	-6.35	-22.88
NBLTM-17×NBLTM-24	1.79	-3.39	-17.39*	-19.67	-26.45	12.31	64.71	48.24	-17.65
NBLTM-17×NBLTM-26	0.00	-6.78*	-20.29*	-42.86	-50.00	-15.38	25.93	25.00	-44.44
NBLTM-17×NBLTM-28	0.85	0.00	-14.49*	-22.33	-27.27	-7.69	-5.00	-17.39	-50.33
NBLTM-17×NBLTM-29	2.80	-6.78*	-20.29*	-55.49	-58.87	-38.46	-17.65	-25.88	-58.82
NBLTM-23×NBLTM-21	5.15	-1.92	-26.09*	10.69	0.51	126.92	114.88	83.33	50.98
NBLTM-23×NBLTM-24	8.16*	0.00	-23.19*	-39.02	-48.89	15.38	12.64	10.11	-35.95
NBLTM-23×NBLTM-26	10.42*	3.92	-23.19*	-55.21	-60.82	-11.54	64.10	43.82	-16.34
NBLTM-23×NBLTM-28	-4.85	15.52*	-28.99*	-1.71	-26.75	65.38	83.43	80.43	8.50
NBLTM-23×NBLTM-29	9.68*	6.65	-26.09*	-20.08	-33.56	50.00	45.98	42.70	-16.99

Table 9 e.Estimates of mid-parent, better parent and standard check heterosis of 25 crosses for seed germination and electrical conductivity in tomato

SCH	Seed germination			Electrical conductivity
SCH	Mid	Better	Standard	Mid	Better	Standard
NBLTM-3×NBLTM-21	5.16*	5.16*	5.16*	-35.15*	-39.64	-39.64*
NBLTM-3×NBLTM-24	36.80*	10.32*	10.32*	-46.63*	-51.79	-51.79*
NBLTM-3×NBLTM-26	10.46*	9.03*	9.03*	-44.52*	-47.66	-47.66*
NBLTM-3×NBLTM-28	-6.40*	-14.81*	3.87*	43.60	15.71*	15.71*
NBLTM-3×NBLTM-29	12.18*	11.46*	12.90*	-22.21	-30.30*	-11.99*
NBLTM-4×NBLTM-21	10.46*	9.03*	9.03*	-55.83	-62.72*	-53.32*
NBLTM-4×NBLTM-24	22.76*	0.00	-2.58*	-28.98	-41.62*	-26.90*
NBLTM-4×NBLTM-26	1.32*	1.32*	-1.29*	-41.27	-49.85*	-37.20*
NBLTM-4×NBLTM-28	2.94*	-7.41*	12.90*	-54.15*	-65.88*	-57.27*
NBLTM-4×NBLTM-29	7.14*	5.10*	6.45*	-69.22*	-69.54*	-61.29*
NBLTM-16×NBLTM-21	26.09*	-6.45*	-6.45*	-58.69*	-59.25*	-63.91*
NBLTM-16×NBLTM-24	68.24*	50.53*	-7.74*	-57.58*	-59.47*	-64.11*
NBLTM-16×NBLTM-26	24.78*	-6.62*	-9.03*	-54.28*	-54.31*	-59.49*
NBLTM-16×NBLTM-28	26.52*	-11.64*	7.74*	-47.19*	-55.36	-60.47*
NBLTM-16×NBLTM-29	28.45*	-5.10*	-3.87*	-57.27*	-63.65*	-54.10*
NBLTM-17×NBLTM-21	24.65*	14.19*	14.19*	-53.44	-61.09	-66.48**
NBLTM-17×NBLTM-24	56.25*	35.66*	12.90*	-36.66*	-45.61	-56.14*
NBLTM-17×NBLTM-26	20.71*	11.92*	9.03*	-25.72*	-38.63	45.59**
NBLTM-17×NBLTM-28	8.81*	-8.47*	11.61*	-27.14*	-29.11	-56.65*
NBLTM-17×NBLTM-29	22.38*	11.46*	12.90*	-46.92*	-61.30	-51.14*
NBLTM-23×NBLTM-21	-6.37*	-7.55*	-5.16*	-48.15*	-50.33	-53.28*

Table 10. Over all heterotic status of crosses in tomato

TESTER	LINES
TESTER	NBLTM-3 (H)	NBLTM-4 (H)	NBLTM-16 (H)	NBLTM-17 (L)	NBLTM-23 (L)
NBLTM-21 (H)	175 (L)	150 (H)	182 (L)	185 (L)	189 (L)
NBLTM-24 (H)	125 (H)	117 (H)	117 (H)	155 (H)	199 (L)
NBLTM-26 (H)	188 (L)	142 (H)	96 (H)	158 (H)	171 (H)
NBLTM-28 (H)	235 (L)	165 (H)	147 (H)	216 (L)	204 (L)
NBLTM-29 (L)	217 (L)	245 (L)	141 (H)	233 (L)	155 (H)

Final norm : 172.28 H: High heterotic status

L: Low heterotic status

Note : Arranged the data in Ascending order

The authors declare potential competing interests as follows:

Determination of general combining ability and Heterosis for yield and quality of tomato inbred lines

Status:

Version 1

Abstract

Figures

Introduction

Materials And Methods

Results And Discussion

Conclusion

Declarations

References

Tables

Additional Declarations

Status:

Version 1