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The butternut squash Cucurbita moschata (Duch. ex Lam.) Duch ex Poir. is an important species for food safety in the world due to its high nutritional value (Restrepo-Salazar et al., 2018a) or for providing medicinal benefits such as improved immune response through β-carotene (Kim et al., 2016). It also presents other medical benefits, such as anti-inflammatory, antioxidant, antidiabetic, antimicrobial, hypotensive, hepatoprotective, antiparasitic, and anticancer properties (Yadav et al., 2010). In addition, this species is used for agro-industrial purposes as food production for humans and animals, and biodiesel production from seed oil (Restrepo et al., 2018b).
Very few studies have been published about the effect of inbreeding on the genetic expression and control of different plant traits in Cucurbita moschata. Espitia (2004) evaluated two diallel crosses of C. moschata (between S0 varieties and between S1 inbred lines) and reported that additive effects played an important role in the expression of the following traits in the two generations evaluated: fruit production per plant, average fruit weight, number of fruits per plant, and 100-seed weight. Non-additive effects were only important in diallel crosses between S1 inbred lines. Similar results were obtained by Ortiz et al. (2013) when evaluating the fruit production per plant in Candelaria (Valle del Cauca, Colombia) and by Restrepo-Salazar et al. (2018a), when evaluating the fruit production per plant and the average fruit weight in the same area. The above-mentioned researchers found, after evaluating three diallel crosses of C. moschata (between S0 parents, between S1 inbred lines, and between S2 lines), that additive effects played an important role in the genetic control of the traits in all inbreeding generations, whereas non-additive effects were only important in crosses between S1 and S2 inbred lines (Ortiz et al., 2013; Restrepo-Salazar et al., 2018a). In other crops, such as maize, records also indicate that non-additive effects are more important in diallel crosses between inbred lines than in crosses between S0 parents (Crossa et al., 1990; Rezende and Souza-Junior, 2000).
According to published literature on diallel crosses of C. moschata, there is no consensus about the type of gene action that predominates in the expression and genetic control of the traits: fruit pulp thickness, number of seeds per fruit, and 100-seed weight. In the case of diallel crosses between S0 parents of C. moschata, some authors reported that both additive and non-additive effects was important in the genetic expression of the fruit pulp thickness (Espitia, 2004; Nisha and Veeraragavathatham, 2014; Abdein et al., 2017), the number of seeds per fruit (Marxmathi et al., 2018; Darrudi et al., 2018), and 100-seed weight (Nisha and Veeraragavathatham, 2014). Other authors like Espitia (2004) and Valdés et al. (2014) found that only additive gene effect was important for 100-seed weight, while Darrudi et al. (2018) reported that only non-additive gene effect was important for the same trait. Espitia (2004), for the number of seeds per fruit, and Marxmathi et al. (2018), for the fruit pulp thickness, found that neither of those effects was important in the expression and genetic control. On the other hand, in the specific case of diallel crosses between S1 inbred lines of C. moschata, studies conducted by Mohanty (2000), Pandey et al. (2010), El-Tahawey et al. (2015), Ahmed et al. (2017), Singh et al. (2018), and Hatwal et al. (2018) reported the importance of both additive and non-additive effects in the expression and genetic control of the fruit pulp thickness. Following the impact of these effects, Mohsin et al. (2017) reported that only the non-additive gene effect was important, whereas other study found that neither of the effects was important (Begum et al., 2016).
According to the background in this field, this study aimed to evaluate the effect of inbreeding on the combining ability for eight traits of butternut squash fruit (C. moschata) and identify parents or F1 hybrids that are outstanding; not only in terms of their combining ability but in terms of the fruit traits.
MATERIALS AND METHODS
Three diallel crosses, each involving six C. moschata parents with three levels of inbreeding (S0 parents, S1 and S2 inbred lines derived from S0 parents), were evaluated at the Experimental Center of the Universidad Nacional de Colombia-Palmira Campus. Table 1 presents the fruit traits of the six S0 parents. A randomized complete block experimental design with four replicates was used. Field treatments were arranged in split plots, with the main plot corresponding to the diallel cross (level of inbreeding), and the subplot was used to evaluate genotypes (six parents and fifteen direct crosses in each of the diallel crosses). Each experimental plot consisted of a five-plant furrow. A weighted selection index composed of traits such as average fruit weight (2.0-4.0 kg), fruit pulp thickness (3.5-5.0 cm) and salmon-colored pulp was used to select the fruits.
The following traits were evaluated: fruit pulp thickness (FPT) measured in cm; fruit pulp color (FPC) ranked from 1 to 15 based on the Roche Yolk Color Fan scale (Vuilleumier, 1969); dry matter (DM) measured as %; diameter of placental cavity (DPC), polar diameter of fruits (PDF), and equatorial diameter of fruits (EDF), all three measured in cm; number of seeds per fruit (NSF); and 100-seed weight measured in g. DM was determined by measuring the fresh weight of fruits and then oven-drying the fruits at 105 °C for 24 hours (Leterme and Estrada, 2012).
Genetic and statistical analyses were performed to estimate the combining ability of the different genotypes, using the method proposed by Hallauer and Miranda (1981), which partitions variation among genotypes (entries) into three components: parents, crosses and the parents vs. crosses contrast. Variance analysis and estimation of genetic effects were performed using the SAS/STAT® package, version 9.4 (SAS system for Windows, SAS Institute Inc©, 2012) and GENES (version 2.1 for Windows©, 2004) developed by Cruz (2013). The F-test was used for several sources of variation during the analysis of variance, and the Student’s t-test was used to estimate genetic effects.
RESULTS AND DISCUSSION
Analysis of variance (ANOVA)
Statistical differences were detected in the source of generational variation for all evaluated traits: FPT, FPC, DM, DPC, PDF, EDF, NSF, and 100-seed weight (Table 2), indicating that at least one of the inbred generations presented a mean value significantly different from the rest. Ortiz et al. (2013) reported similar results in C. moschata for FPC. On the other hand, significant differences were observed in the source of genotype variation in the three generations analyzed for all the traits. It can be inferred that there is at least one parent or hybrid that recorded an average value of FPT, FPC, DM, DPC, PDF, EDF, NSF, and 100-seed weight differed statistically from other averages in each of the generations (Table 2). Espitia (2004) also recorded statistical differences in the source of C. moschata genotypes for FPT, NSF, and 100-seed weight in the two inbreeding generations studied (S0 and S1). Ortiz et al. (2013) observed similar results in C. moschata for FPC, finding differences among genotypes in the three diallel crosses evaluated (S0, S1, and S2).
Table 2 Mean squares of ANOVA for fruit pulp thickness (FPT), fruit pulp color (FPC), dry matter (DM), diameter of placental cavity (DPC), polar diameter of fruits (PDF), equatorial diameter of fruits (EDF), number of seeds per fruit (NSF), and 100-seed weight in three inbred generations of Cucurbita moschata.
For all evaluated traits, in most of the cases the parents and crosses variation sources presented statistical significance in all three generations, which confirmed that in general terms at least one of the S0, S1, or S2 parents, or at least one of the crosses between said parents, showed an average performance of FPT, FPC, DM, DPC, PDF, EDF, NSF, and 100-seed weight that differed significantly from the others (Table 2). Based on these results, it is inferred that regardless of the inbreeding level, it is possible to identify at least one parent or a hybrid with a mean value, in any of the traits, that differs statistically from the others. Significant differences have also been reported in C. moschata by Espitia (2004) regarding FPT, NSF, and 100-seed weight as well as by Ortiz et al. (2013) regarding FPC in both parents and crosses as sources in the different inbred generations under study.
In the source of variation corresponding to the contrast between parents vs. crosses (P vs. C), significant differences were detected for most of the traits of the inbred generations, S1 (FPT, FPC, DPC, PDF, EDF, NSF, 100-seed weight) and S2 (FPT, DPC, PDF, EDF, NSF, 100 seed-weight), indicating that, overall, the average performance of all F1 crosses (between S1 or S2 inbred lines) was higher than the average performance of parents as a whole (Table 2). Similar results were found by Espitia (2004) in S1 for FPT, NSF, and 100-seed weight. On the other hand, in the S0 inbred generation, statistical differences were only observed in the P vs. C contrast for DPC, EDF, NSF, and 100-seed weight (Table 2). Espitia (2004) reported similar results in S0 for NSF but did not record differences for 100-seed weight in the S0 generation in the contrast P vs. C.
In the diallel cross between S0 parents only, the additive effects (GCA) were important in the genetic expression and control of FPT (Table 2). In contrast, other authors have reported in S0 parents that in C. moschata both additive and non-additive effects (SCA) are important in the genetic expression of FPT (Espitia, 2004; Nisha and Veeraragavathatham, 2014; Abdein et al., 2017), while other authors have published that neither of the effects was important for FPT (Marxmathi et al., 2018). Regarding diallel crosses between S1 and S2 inbred lines, results indicated the importance of the additive and non-additive gene effect in the genetic control of FPT (Table 2). Mohanty (2000), Pandey et al. (2010), El-Tahawey et al. (2015), Ahmed et al. (2017), Singh et al. (2018), and Hatwal et al. (2018) reported similar results in the cross between S1 inbred lines of C. moschata. In contrast, a study conducted by Begum et al. (2016) involving crosses between S1 inbred lines of C. moschata indicate that neither of the effects was important in the expression of FPT, whereas Mohsin et al. (2017) only reported the importance of non-additive effects.
In the diallel cross between S0 parents, neither of the two types of effects was important in the genetic expression and control of NSF (Table 2). This result indicates, on the one hand, that there is not enough statistical evidence to conclude that some parents differ in its ability to transmit genes that allow its progeny to increase or decrease its NSF; on the other hand, it suggests that there is not enough evidence to conclude that some of the hybrids had a different behavior than expected based on combining ability of their parents and general mean. Espitia (2004) reported similar results in C. moschata, finding that the additive and non-additive effects did not contribute in statistical terms to the genetic expression of NSF. In contrast, Marxmathi et al. (2018) and Darrudi et al. (2018) found that both types of effects were important for NSF. The importance of additive and non-additive gene action in the control of NSF was observed in the case of diallel crosses between S1 and S2 inbred lines (Table 2). Espitia (2004), El-Tahawey et al. (2015), and Mohsin et al. (2017) also found a significant contribution of both types of effects on the genetic expression of NSF in diallel crosses between S1 inbred lines of C. moschata.
Both additive and non-additive effects were important in the genetic control of the 100-seed weight in the three diallel crosses evaluated (Table 2). These results agree with those found by Nisha and Veeraragavathatham (2014) in diallel crosses of C. moschata between S0 parents, and by Espitia (2004), Mohsin et al. (2017) and Hatwal et al. (2018) in diallel crosses between S1 inbred lines. On the other hand, Espitia (2004) reported that only additive effects were important in the expression of 100-seed weight in diallel crosses between S0 parents of C. moschata, while Darrudi et al. (2018) reported that only non-additive effects were important. Valdés et al. (2014) recorded a differential response in diallel crosses between S0 parents of C. moschata evaluated during two different planting seasons, finding that both additive and non-additive effects were important in the genetic expression of 100-seed weight during one season, while only additive effects were responsible for its expression in the same genotypes during another season.
In the diallel cross between S0 parents, only additive effects were important in the genetic expression and control of PDF and EDF (Table 2). In contrast, Abdein et al. (2017), Kakamari and Jagadeesha (2017), and Marxmathi et al. (2018) have reported that both additive and non-additive effects are important in the genetic expression of PDF and EDF in C. moschata. In the case of diallel crosses between S1 inbred lines, this study indicates that both types of effects were important in the genetic control of both traits. These results are similar to those reported by Jha et al. (2009), Ahmed et al. (2017), Mohsin et al. (2017), and Singh et al. (2018) for diallel crosses between S1 parents in C. moschata and by Rana et al. (2015) in diallel crosses between advanced inbred lines.
Additive effects were the only component of important variation in the genetic control of DPC and DM in the diallel cross between S0 parents (Table 2). Marxmathi et al. (2018), on the other hand, reported that no effect was important for DM. Both additive and non-additive gene actions were observed to be important in the genetic control of DPC and DM in diallel crosses between S1 and S2 inbred lines (Table 2). Similar results were recorded by Rana et al. (2015) for DPC in diallel crosses between advanced lines of C. moschata; both types of effects were found to control these traits. Regarding DM, these same authors reported that only additive effects contributed significantly to its genetic expression. On the other hand, the analysis of data for FPC indicated that additive effects were important in its genetic expression and control only in diallel crosses between S0 parents, whereas non-additive effects were important in its control in diallel crosses between S0 parents and between S2 inbred lines (Table 2). In contrast, Ortiz et al. (2013) reported the importance of additive effects in all generations evaluated (S0, S1, S2), while non-additive effects were only important in the expression of FPC in the S0 generation.
The joint analysis of the results of the three diallel crosses indicates that, in general terms, additive effects were responsible for the genetic expression and control for most of the traits in crosses made between the different inbred generations evaluated. Non-additive effects, on the other hand, were also responsible for the genetic control for most of the traits, but almost exclusively in the crosses made between S1 and S2 inbred lines. This could be attributed to the greater genetic divergence occurring in crosses between parents with a narrow genetic base in contrast to crosses between broad-based parents. Espitia (2004) observed similar results in C. moschata for 100-seed weight, yield, and yield components in diallel crosses between S0 parents and between S1 lines.
General combining ability (GCA) effects
A differential response was observed in parents in their general combining ability (GCA) effects for FPT, indicating the highly significant differences in additive effects detected by ANOVA in the three diallel crosses (Table 2). The S0 parents (P3 and P4) and the S2 inbred lines (P1, P2, and P3) presented significant GCA effects values as well as highest FPT values (Table 3). Espitia (2004), Nisha and Veeraragavathatham (2014), and Abdein et al. (2017) also recorded at least one S0 parent with significant GCA effects values. Other authors (Mohanty, 2000; Espitia, 2004; Pandey et al., 2010; El-Tahawey et al.; 2015, Ahmed et al., 2017; Singh et al., 2018; Hatwal et al., 2018) have also reported the existence of at least one S1 line with significant GCA effects values. However, Rana et al. (2015) and Begum et al. (2016) did not find any inbred line of C. moschata with significant GCA effects values. Of the outstanding genotypes mentioned previously, S0 parents (P3 and P4) are recommended to genetically improve FPT for the fresh consumption market formed by consumers who prefer whole, non-sliced fruits, taking advantage of the additive effects of intrapopulation recurrent selection (IRS). The S2 inbred line (P2) is recommended for the improvement of FPT for agro-industrial use or the fresh consumption market formed by consumers for whom fruit weight is not a limiting characteristic for purchase (Table 3).
Table 3 Cucurbita moschata parents showing general combining ability (GCA) effects that are significant for fruit pulp thickness (FPT), fruit pulp color (FPC), polar diameter of fruits (PDF), and 100-seed weight, obtained in diallel crosses between S0 parents and between S1 and S2 inbred lines.
When evaluating FPC, almost all the S0 parents and S1 and S2 inbred lines recorded significant GCA effects values and a medium orange color according to the Roche Yolk Color Fan scale (Vuilleumier, 1969) (Table 3). Of these genotypes, S0 parents (P3 and P4) are recommended for the genetic improvement of FPC destined for the fresh consumption market. This market is formed by consumers who prefer whole, non-sliced, fruits. The S2 inbred line (P2) is recommended for agro-industrial use or the fresh consumption market formed by consumers for whom fruit weight is not a limiting characteristic for purchase. Ortiz et al. (2013) also identified in C. moschata at least one S0 parent or S1 and S2 lines with significant GCA effects values for FPC.
In the case of 100-seed weight, the only genotypes that reported significant GCA effects values were the S1 inbred line (P2) and the S2 inbred line (P5) (Table 3). These genotypes, however, did not show high 100-seed weight values. Other authors haver reported similar results for 100-seed weight in S1 lines (Espitia, 2004; Mohsin, 2017; Hatwal et al., 2018). Espitia (2004) found at least one S0 parent of C. moschata with significant GCA effects values for 100-seed weight, whereas Valdés et al. (2014) did not find any S0 parent with significant GCA effects values.
Only the S2 line (P5) recorded significant GCA effects value for PDF (Table 3). Similar results have been reported in C. moschata by several authors (Jha et al., 2009; Rana et al., 2015; Ahmed et al., 2017; Mohsin et al., 2017; Singh et al., 2018), who found at least one inbred line with significant GCA effects values, indicating that for this specific trait, only a few genotypes evaluated had the ability to transmit favorable genes to their progenies. Kakamari and Jagadeesha (2017) and Marxmathi et al. (2018) reported at least one S0 parent with significant GCA effects values for PDF. Finally, in the case of DM, DPC, EDF, and NSF, no parent was genetically superior to the other parents under this study (Table 3). Rana et al. (2015) did not find any inbred line of C. moschata with significant GCA effects values for DPC and EDF; however, they did record at least one inbred line with a significant GCA effects value for DM. Jha et al. (2009) also reported the non-existence of C. moschata lines with significant GCA effects values for EDF. However, Ahmed et al. (2017), Mohsin et al. (2017), and Singh et al. (2018) recorded significant GCA effects for this trait in inbred lines, whereas Kakamari and Jagadeesha (2017), and Marxmathi et al. (2018) reported at least one S0 parent with significant GCA effects values for EDF. Furthermore, other authors (Espitia, 2004; El-Tahawey et al., 2015; Mohsin et al., 2017) identified at least one inbred line of C. moschata with significant GCA effects values for NSF, differing from the results found in the current study.
The joint analysis of all the traits evaluated in this study indicated that, for butternut squash destined to the fresh consumption market formed by consumers who prefer whole, non-sliced, fruits, S0 parents (P3 and P4) can be suggested as genotypes to improve FPT and FPC genetically, taking advantage of the additive effects of IRS. These parents presented significant GCA effects values for both traits, with P3 presenting an FPT of 4.40 cm and P4, one of 4.30 cm; both presented a medium orange FPC (Table 3). In addition, they presented acceptable average values for the other studied traits. On the other hand, in the case of butternut squash for agro-industrial use or the fresh consumption market consisting of consumers for whom fruit weight is not a limiting characteristic for purchase, S2 parent (P2) can be recommended for the simultaneous improvement of FPT and FPC, taking advantage of both additive and non-additive effects by IRS. This genotype reported significant GCA effects values for both traits, with an FPT of 4.70 cm and a medium orange FPC (Table 3). It also presented acceptable average values for the other traits under study.
Specific combining ability (SCA) effects
Several crosses between S0 parents or between S1 and S2 inbred lines presented significant SCA effects for FPT (Table 4), presenting values above the expected average based on the GCA effects values of parents and the overall average. Similar results were observed in C. moschata by Espitia (2004), who reported at least one cross between S0 parents with significant SCA effects values. Other authors have also reported the existence of at least one inbred line with significant SCA effects values for FPT in C. moschata (Mohanty, 2000; Pandey et al., 2010; El-Tahawey et al., 2015; Rana et al., 2015; Ahmed et al., 2017; Mohsin et al., 2017; Singh et al. 2018; Hatwal et al., 2018). However, Espitia (2004) and Begum et al. (2016) did not find any S1 inbred line of C. moschata with significant SCA effects values for FPT. Of the outstanding hybrids mentioned; in the case of the fresh consumption market formed by consumers who prefer whole, non-sliced fruit. The hybrids between S0 parents (P1×P4) and (P2×P3) are the genotypes recommended to improve FPT, taking advantage of additive effects and allowing superior-performance varieties or lines obtained by transgressive segregation. The S2 hybrid (P1×P6) is recommended to improve FPT for the same market, using reciprocal recurrent selection (RRS) to take advantage of both types of effects. In the case of genotypes for agricultural use or for the fresh consumption market formed by consumers for whom fruit weight is not a limiting characteristic for purchase, the between S1 inbred line hybrids (P1×P5), the S2 inbred line hybrids (P1×P3), and (P2×P6) are recommended for improving FPT by RRS (Table 4).
Table 4 Hybrids of Cucurbita moschata obtained by diallel crosses between S0 parents and between S1 and S2 inbred lines, showing significant combining ability effects (SCA effects) for fruit pulp thickness (FPT), fruit pulp color (FPC), dry matter (DM), diameter of placental cavity (DPC), polar diameter of fruits (PDF), equatorial diameter of fruits (EDF) and 100-seed weight.
The analysis of FPC indicated that several crosses between S0 parents or between S1 and S2 inbred lines were identified with significant SCA effects values (Table 4). Of these crosses, the hybrid between S2 inbred line (P1×P6) is recommended to genetically improve FPC for the fresh consumption market formed by consumers who prefer whole, non-sliced fruits. Ortiz et al. (2013) had also reported the existence of at least one cross between S0 parents in C. moschata with a significant SCA effects value.
In the case of DPC, three hybrids between S0 parents and two between S2 inbred lines presented significant SCA effects values (Table 4). Of these, the hybrid S2 inbred line (P2×P6) is recommended to improve DPC by RRS for agro-industrial use or for the fresh consumption market formed by consumers for whom fruit weight is not a limiting characteristic for purchase. Rana et al. (2015) also reported the existence of at least one hybrid between advanced inbred lines of C. moschata with significant SCA effects values for DPC.
In the case of 100-seed weight, two hybrids between S0 parents, one between S1 inbred lines and another between S2 inbred lines showed significant SCA effects values (Table 4). Similar results were reported in C. moschata by several authors (Espitia, 2004; Valdés et al., 2014; Nisha and Veeraragavathatham, 2014), who observed at least one cross between S0 parents with significant SCA effects values. Other authors have also reported the existence of at least one inbred line of C. moschata with significant SCA effects values for 100-seed weight (Espitia, 2004; El-Tahawey et al., 2015; Mohsin et al., 2017; Hatwal et al., 2018).
One hybrid between S0 parents and another between S1 inbred lines presented significant SCA effects values for DM (-1.29* and 2.19*, respectively) (Table 4), indicating that their DM contents were lower in the first hybrid and higher in the second one, with respect to expected mean with base in GCA effects of its parents and general mean. Rana et al. (2015) had also observed the existence of at least one hybrid between advanced inbred lines of C. moschata with significant SCA effects values for DM.
Only two crosses with significant SCA effects values were identified in the case of EDF. These corresponded to hybrids between the S0 parents (P2×P6) and (P5×P6) (Table 4). Kakamari and Jagadeesha (2017) and Marxmathi et al. (2018) found similar results in hybrids between S0 parents for EDF. On the other hand, the existence of at least one hybrid between inbred lines of C. moschata, with significant SCA effects values for EDF, has been reported by other authors (Jha et al., 2009; Rana et al., 2015; Ahmed et al., 2017; Mohsin et al., 2017; Singh et al., 2018).
This study only revealed one cross with a significant SCA effects value for PDF, the hybrid between S2 inbred lines (P5×P6) (Table 4). Jha et al. (2009), Ahmed et al. (2017), Mohsin et al. (2017) and Singh et al. (2018) had also recorded the presence of at least one hybrid between inbred lines with significant SCA effects values for PDF. Kakamari and Jagadeesha (2017) and Marxmathi et al. (2018), on the other hand, reported at least one hybrid between S0 parents with significant SCA effects values for PDF.
No crosses presented significant SCA effects values for NSF. Espitia (2004) had not reported the existence in C. moschata of crosses between S1 inbred lines with significant SCA effects values. However, the presence of at least one cross between inbred lines with significant SCA effects values has been observed by other authors (El-Tahawey et al., 2015; Mohsin et al., 2017). Espitia (2004), Marxmathi et al. (2018), and Darrudi et al. (2018), on the other hand, reported at least one cross between S0 parents with significant SCA effects values for NSF.
The joint evaluation these results indicated that in the case of the fresh consumption market formed by consumers who prefer whole, non-sliced fruits; the hybrid between S2 inbred lines (P1×P6) is the cross recommended for the simultaneous genetic improvement of FPT and FPC. It takes advantage of both additive and non-additive effects by RRS and records a significant SCA effects value for both traits, presenting an FPT of 4.01 cm and a medium orange FPC (Table 4). In addition, this hybrid presented acceptable average values for the other traits studied.
CONCLUSIONS
Additive effects were responsible for the genetic expression and control for most of the fruit traits evaluated in the three diallel crosses. Non-additive effects were also responsible for the genetic control of most of the traits, but almost exclusively in crosses between S1 and S2 inbred lines. In the case of the fresh consumption market formed by consumers who prefer whole, non-sliced fruits, the S0 parents UNAPAL-Dorado and IC3A (P3 and P4) were recommended for the simultaneous genetic improvement of traits fruit pulp thickness and fruit pulp color, taking advantage of additive effects. In the case of genotypes destined for agro-industrial use or for the fresh consumption market formed by consumers for whom fruit weight is not a limiting characteristic for purchase, the S2 parent UNAPAL-Abanico-75-2 (P2) is recommended for the simultaneous improvement of traits fruit pulp thickness and fruit pulp color, taking advantage of additive effects. In the case of the fresh consumption market formed by consumers who prefer whole, non-sliced fruits, the hybrid between S2 inbred lines UNAPAL-Abanico-75-1×UNAPAL-Llanogrande-2 (P1×P6) is recommended for the simultaneous genetic improvement of traits FPT and FPC, taking advantage of both additive and non-additive effects. After evaluating the effect of inbreeding on the genetic expression and control of fruit traits analyzed in this study, it was found that non-additive effects are important in diallel crosses between inbred lines than in those between S0 parents.
