Open-access Genetic mean components as selection criteria for identifying promising yellow melon hybrids for inbred line extraction

Componentes de média genética como critérios de seleção para identificação de híbridos de melão amarelo promissores para extração de linhagens endogâmicas

Abstract

Estimates of the mean components m + a′ and d represent a promising strategy for guiding the selection of populations and lines in plant breeding programs. This study estimated m + a′ and d to identify commercial yellow melon hybrids suitable for inbred line extraction to develop new hybrids. The experiments evaluated seven commercial single-cross (F1) yellow melon hybrids and their self-pollinated progenies (F2), for a total of 14 populations. Experiments were conducted in a randomized complete block design with three replications in a split-plot arrangement, with generation assigned to main plots and hybrids to subplots. Evaluated traits included fruit number per plot, mean fruit mass, yield, flesh thickness, flesh firmness, and total soluble solids. The Mulamba and Mock selection index (MMI) provided joint selection across all traits. Individual and combined analyses of variance applied a split-plot model and treated generation and hybrid effects as fixed. In the F1 generation, homozygous gene effects (m + a′) predominated and contributed substantially to all traits and the MMI. Estimates of m + a′ and d classified hybrids by success probability in inbred line extraction and showed that inbreeding depression was not significant for all melon traits. Hybrids AF-13395 and Goldmine showed the highest suitability for selecting superior inbred lines.

Keywords:
Cucumis melo L.; plant breeding; homozygosity; genetic mean components; inbreeding depression

Resumo

As estimativas dos componentes de média m + a′ e d fornecem um método para seleção de populações e linhagens em programas de melhoramento genético vegetal. Este estudo estimou

m + a′ e d para identificar híbridos comerciais de melão amarelo adequados para extração de linhagens endogâmicas visando o desenvolvimento de novos híbridos. Os experimentos avaliaram sete híbridos comerciais de cruzamento simples (F1) de melão amarelo e suas respectivas progênies autopolinizadas (F2), totalizando 14 populações. Os experimentos utilizaram delineamento de blocos casualizados completos com três repetições em arranjo de parcelas subdivididas, com gerações alocadas nas parcelas principais e híbridos nas subparcelas. As características avaliadas incluíram número de frutos por parcela, massa média de frutos, produtividade, espessura de polpa, firmeza de polpa e sólidos solúveis totais. O índice de seleção de Mulamba e Mock (MMI) permitiu seleção conjunta de todas as características. Análises individuais e combinadas de variância aplicaram modelo de parcelas subdivididas, considerando efeitos de geração e híbrido como fixos. Na geração F1, efeitos gênicos homozigotos (m + a′) predominaram e contribuíram substancialmente para todas as características e para o MMI. As estimativas de m + a′ e d classificaram os híbridos pela probabilidade de sucesso na extração de linhagens consanguíneas e demonstraram que a depressão por endogamia não foi significativa para todas as características avaliadas em melão. Os híbridos AF-13395 e Goldmine apresentaram maior adequação para seleção de linhagens endogâmicas superiores.

Palavras-chave:
Cucumis melo L.; melhoramento genético vegetal; homozigose; componentes de média genética; depressão por endogamia

1. Introduction

Melon (Cucumis melo L.) ranks among the fruit crops most widely cultivated and consumed worldwide (Cui et al., 2022) and constitutes a primary cucurbit crop in the Brazilian Semiarid region. In Brazil, melon production concentrates in the states of Rio Grande do Norte, Bahia, and Ceará, which account for more than 95% of national production (IBGE, 2024).

Sustaining and expanding melon production in the Northeast region of Brazil requires ongoing breeding to develop new hybrids that exceed current ones. These efforts depend on quantifying genetic components of hybrid performance. The mean value of any trait in a single-cross hybrid (HS) can be expressed as HS = m + a′ + d, where m + a′ represents the contribution of homozygous loci already fixed in the parental inbred lines (corresponding to the mean of the two parents), and d denotes the contribution of heterozygous loci, commonly referred to as heterosis or hybrid vigor (Bernardo, 2020; Ramalho et al., 2024). Superior hybrid development requires productive inbred lines and dominance effects from their crosses.

Extracting high-performing inbred lines requires identifying source populations with high success probability in breeding. Commercial hybrids, subjected to intensive selection for adaptation and performance in regional conditions, serve as genetic sources for extracting inbred lines. In melon breeding, researchers have used commercial hybrids from seed companies to identify populations suitable for inbred line extraction. This practice aligns with the Brazilian Plant Variety Protection Law (Law No. 9,456/1997), which allows inbred line extraction from protected commercial hybrids (Brasil, 1997).

Estimates of the mean components m + a′ and d from commercial hybrids identify those suitable for inbred line development. Brazilian melon hybrids consist mainly of single crosses from two parental lines, with segregating locus frequency of 0.5. Under these conditions, the m + a′ estimate represents the expected mean of inbred lines after complete inbreeding (F∞), without selection. This parameter quantifies segregating population potential from fixed loci in parental lines.

The d estimate quantifies contributions of heterozygous loci and expected genetic variability across inbreeding generations. Higher d values indicate greater heterozygosity in the hybrid and superior inbred lines via selection, as favorable alleles fix in homozygosity and join fixed alleles (Ramalho et al., 2024).

Researchers apply mean component estimates to select populations in breeding programs for self-pollinated crops, including common bean (Phaseolus vulgaris) (Oliveira et al., 1996; Abreu et al., 2002; Carneiro et al., 2002; Mendonça et al., 2002; Rocha et al., 2015) and cowpea (Vigna unguiculata) (Matos et al., 2020). In maize (Zea mays), this method applies to commercial hybrids (Lima et al., 2000; Souza Sobrinho, 2001; Viana et al., 2009). No studies report mean component estimates across multiple commercial melon hybrids.

Most studies on m + a′ and d estimates examine yield due to its economic value in major crops. In melon breeding, yield and fruit quality traits—soluble solids, flesh thickness, and flesh firmness—require joint consideration for market acceptance (Oliveira et al., 2019). Melon fruit, as a consumer product, necessitates the evaluation of yield and quality traits together when selecting populations for inbred line extraction.

This study estimated homozygous (m + a′) and heterozygous (d) locus contributions to agronomic traits in commercial melon (Cucumis melo L.) hybrids to identify those suitable for inbred line extraction and hybrids adapted to the Northeast region of Brazil.

2. Material and Methods

2.1. Experimental conditions

Two experiments were conducted: Experiment 1 from September to November 2017 in Baraúna (5°04′S, 37°37′W; 109 m altitude) and Experiment 2 from August to October 2018 in Alagoinha, Mossoró, Rio Grande do Norte, Brazil (5°03′S, 37°24′W; 70 m altitude). The regional climate is classified as BSh, hot semiarid, according to the Köppen-Geiger classification, with a dry season from June to January and a rainy season from February to May (Alvares et al., 2013). Crop management practices followed recommendations for Rio Grande do Norte, Brazil (Nunes et al., 2016).

2.2. Plant material and experimental design

Seven single-cross commercial yellow melon hybrids (F1) were evaluated: Iracema, AF-13395, Goldex, Gladial, Goldmine, Mandacaru. and Solares. Self-pollinated progenies (F2) yielded 14 populations total. Experiments were conducted in a randomized complete block design with three replications in a split-plot arrangement, with generation assigned to main plots and hybrids to subplots. Each plot had seven plants in a single row, spaced 0.4 m apart. Over two years, F1 and F2 populations of the seven hybrids formed 14 treatments. Harvest samples of five commercial fruits per plot were evaluated.

2.3. Traits

The agronomic traits evaluated were fruit number per plot, mean fruit mass, yield, flesh thickness, flesh firmness, and soluble solids. Fruit number per plot was determined by a total fruit count. Mean fruit mass (g) was calculated as the average mass of five fruits per plot. Yield (Mg ha−1) was determined by summing the mass all fruits harvested in the plot; fruits were weighed individually on an electronic scale. Flesh thickness (mm) was measured as the mean pericarp thickness of five fruits per plot (two measurements per fruit), using a graduated scale. Flesh firmness (kgf) was measured by two penetration measurements per side of the equatorial flesh region after a transverse cut, using a Wagner penetrometer equipped with an 8.0 mm conical plunger (FT 011 Fruit Pressure Tester). Soluble solids content (°Brix) was determined as the mean from two equatorial flesh pieces (lower two-thirds) of five fruits per plot, pressed into a Palette 100 digital refractometer. The samples were manually pressed directly into a digital refractometer (Palette 100), and results were expressed as the mean of five fruits per plot.

2.4. Statistical analysis

Individual and joint analyses of variance applied a split-plot model to all traits, with generation and hybrid as fixed effects and year as random. The Mulamba and Mock (1978) selection index (MMI) enabled joint multi-trait selection. MMI weights (w) followed farmer input on trait importance: fruit number (w1 = 0.15); mean fruit mass (w2 = 0.15); yield (w3 = 0.30); flesh thickness (w4 = 0.10); flesh firmness (w5 = 0.10); and soluble solids (w6 = 0.20).

The contributions of homozygous and heterozygous loci were estimated using the generation mean components F1=m+a+d and F2=m+a+12d, where m is the general mean, a′ is the additive effect of homozygous loci, and d is the dominance effect of heterozygous loci. From these expressions, the homozygous contribution (m + a′) was calculated as 2F¯2F¯1, and the heterozygous contribution (d) as 2F¯1 F¯2. The contributions of homozygous and heterozygous loci were expressed as percentages. The significance of these parameters was assessed via Student’s t-test. Analysis of variance (ANOVA) was performed using the SISVAR® 5.6 software (Ferreira, 2019). Alls analyses used a significance level of α = 0.05.

3. Results

3.1. ANOVA and inbreeding estimates

Generations differed significantly for fruit number, yield, flesh thickness, and Mulamba-Mock index (MMI) (Table 1). Hybrids differed for all traits (Table 1). The effect of the hybrid × year interaction was significant for fruit number and soluble solids, indicating hybrid rank changes across years. The effect of the hybrid × generation × year interaction was significant for soluble solids (Table 1), with performance varying across generation-year combinations.

Table 1
Snedecor's F-values, overall means, mean square errors, and coefficients of variation from analysis of variance for six traits and Mulamba-Mock index (MMI) in F1 and F2 generations of yellow melon hybrids across two years in Baraúna and Mossoró, Rio Grande do Norte, Brazil, 2017–2018.

Hybrids showed similar means for mean fruit mass, flesh thickness, flesh firmness, and soluble solids across F1 and F2 (Table 2). F2 means for all hybrids fell within the export range for yellow melon fruit, indicating that both F1 and F2 generations supported commercialization. Yield means declined from F1 to F2 for most hybrids (Table 2). Some hybrids yielded below 25 Mg ha−1, the farmer-defined minimum yield to ensure profitability. MMI showed low variation among hybrids, particularly in F2. F1 exceeded F2 for fruit number, yield, and flesh thickness (Table 2).

Table 2
Mean of agronomic traits of the F1 and F2 generations of yellow melon hybrids evaluated in two consecutive years in Baraúna and Mossoró, Rio Grande do Norte, Brazil, 2017–2018.

Inbreeding depression estimates and Mulamba-Mock index (MMI) are shown in Table 3. Inbreeding depression magnitude varied among traits and MMI, with the highest estimates for yield and the lowest for soluble solids. Hybrids varied in inbreeding depression for all traits and MMI. Yield inbreeding depression ranged from 44.84% to -15.80% (60.64 percentage points total range) (Table 3). Soluble solids inbreeding depression ranged from -0.17% to 0.36% (Table 3). MMI showed the highest mean inbreeding depression.

Table 3
Inbreeding depression estimates for agronomic traits of yellow melon hybrids across two years in Baraúna and Mossoró, Rio Grande do Norte, Brazil, 2017–2018.

3.2. Estimates of m + a′ and d

Homozygous loci (m + a′) and heterozygous loci (d) contributions to the six traits are shown in Table 4. Student's t-test indicated significant m + a′ for all hybrids and traits. Hybrid Mandacaru exhibited the highest m + a′ estimates for fruit number and yield, whereas hybrid Iracema showed the lowest. The hybrid Gladial exhibited the highest m + a′ value for mean fruit mass and flesh thickness, whereas hybrid Iracema showed the lowest. Hybrids AF-13395 exhibited the highest m + a′ for flesh firmness, whereas hybrid Solares showed the lowest. The m + a′ estimates for soluble solids varied little among hybrids, with hybrid AF-13395 showing the highest value. The heterozygous loci (d) contribution was significant for yield (all hybrids) and fruit number (all hybrids except Gladial). Hybrid AF-13395 exhibited the highest yield d, whereas Gladial showed the lowest (Table 4).

Table 4
Homozygous locus (m + a′) and heterozygous locus (d) contributions to six traits and MMI of yellow melon hybrids across two years in Baraúna and Mossoró, Rio Grande do Norte, Brazil, 2017–2018.

The mean m + a′ of the seven hybrids exceeded d means, indicating greater additive effects from parental lines than F1 dominance effects. The hybrid m + a′ mean percentages exceeded d mean percentages in 80% for five traits, from 67.08% (mean fruit mass) to 99.65% (soluble solids) (Figure 1). These results suggest that additive genetic effects predominate over dominance effects for melon traits.

Figure 1
Percentage of homozygous loci (m + a′) and heterozygous loci (d) to six traits and MMI of yellow melon hybrids across two years in Baraúna and Mossoró, Rio Grande do Norte, Brazil, 2017–2018. FN = fruit number per plot; AFW = mean fruit mass (kg); YID = yield (Mg ha−1); FT = flesh thickness (cm); FF = flesh firmness (kgf); SS = soluble solids (°Brix); MMI = Mulamba-Mock index.

4. Discussion

Genotype evaluation requires high experimental precision, as cultivar differences are small in most crops. High precision improves the detection of differences among treatments (Ramalho et al., 2000). Selective accuracy (As) quantifies experiment quality. This study recorded high selective accuracy (As > 0.90) for all traits, confirming high experimental quality (Resende and Alves, 2022). The coefficient of variation (CV) provides the standard experimental quality measure. According to the classification of Lima et al. (2004), CV estimates for melon yield and total soluble solids content were considered average (Table 1). The CV values are consistent with those observed in the Mossoró–Jaguaribe hub experiments (Oliveira et al., 2019; Cavalcante Neto et al., 2025; Lima et al., 2025).

Hybrid effect significance showed genotypic variation for all traits (Table 1). Generations differed for fruit number, yield, and flesh thickness. F1 means exceeded F2 means for these traits, confirming inbreeding depression (Table 3). Inbreeding depression reflects genotypic value reduction from increased homozygosity, decreased heterozygosity, and dominance interactions.

Inbreeding depression varied among traits (Table 3). Trait-specific heritability, numbers of genes (loci), allelic interactions, and gene interactions (epistasis) explained this variation. Quantitative trait inheritance amplified environmental influence on phenotypic expression. Hybrid variation in inbreeding depression resulted from parental allele complementarity (genetic divergence) and dominance or epistasis (non-additive effect) (Table 3). Limited parental allele complementarity or low dominance (non-additive effect) results in low inbreeding depression, whereas greater parental allele complementarity increases inbreeding depression and reduces heterosis (Charlesworth and Willis, 2009).

Classical models explain inbreeding depression and heterosis via dominance and overdominance, the literature also describes the effects additive × additive epistasis, which may result in negative heterosis (Lamkey and Edwards, 1999; Bernardo, 2020; Ramalho et al., 2024) or pseudo-dominance (Wallace et al., 2014). Models may operate simultaneously across allogamous and autogamous crops, with inbreeding depression and heterosis varying by genotype.

Cucurbitaceae species show low inbreeding depression despite cross-pollination favored by floral structure. Evolutionary processes of these species reduced inbreeding effects, as plants cover large areas with few individuals, promoting related mating (inbreeding), and natural selection eliminate deleterious alleles over time (Allard, 1999). Low inbreeding depression has been observed in pumpkin (Cardoso, 2004), watermelon (Wehner, 2008), and melon (Monforte et al., 2004).

Studies that guide selection of populations and lines increase breeding program efficiency because evaluating segregating populations is costly and time-consuming. Traditionally, breeders generate segregating populations with superior agronomic traits using diallel crosses, an effective method in several crops (Pouyesh et al., 2017; Cavalcante Neto et al., 2020; Napolitano et al., 2020; Shajari et al., 2021; Melgoza et al., 2022; Rad et al., 2023; Soltani et al., 2023). However, diallel crosses demand substantial time and labor, increasing breeding program cost. Therefore, alternative strategies are needed to manage segregating populations and identify superior lines in early generations.

Estimating m + a′ and d provides an alternative to identify superior populations early and efficiently. The m + a′ estimate identifies populations with higher means when all loci are homozygous, assuming no selection. The d estimate indicates dominance in genetic control of the trait. Higher d values imply more loci in heterozygosity and therefore greater genetic variability expressed in subsequent inbreeding generations. If selection is efficient, additional loci with favorable alleles increase mean inbred line performance. Thus, d functions as an indicator of potential population variability. Breeders should select populations with high m + a′ and high d to develop new inbred lines. Abreu et al. (2002) reported this pattern in common bean, where m + a′ estimates from F1 and F2 efficiently predicted seed yield and variance among F7 populations.

Estimates of m + a′ varied among traits (Table 4), indicating fixed homozygous loci for these traits. Populations with higher m + a′ show greater frequency of fixed favorable alleles at allele frequency of 0.5 (Viana et al., 2009; Rocha et al., 2015; Ramalho et al., 2024). These m + a′ and d estimates indicate low heterosis in melon. However, yield showed heterosis in some hybrids. Yield and soluble solids represent primary traits in melon breeding (Oliveira et al., 2019). Single-cross hybrids remain viable in melon despite low heterosis and inbreeding depression, as Brazilian farmers have used them since the mid-1980s in major production regions. Fruit uniformity and resistance to pathogens and insect pests further support the use of simple hybrids.

Prior studies estimated these parameters in allogamous species (e.g., maize) and autogamous species (e.g., cowpea). Maize single-cross hybrids showed higher d than m + a′ for grain yield (Lima et al., 2000). Contrastingly, cowpea yield showed predominant m + a′ (Matos et al., 2020). Thus, genetic interactions vary by species, requiring crop-specific analysis.

Previous studies emphasized grain yield as the primary selection trait in major crops. However, melon breeding requires consideration of fruit yield and quality traits. Thus, the present study evaluated the primary traits identified by melon farms in the Mossoró–Jaguaribe hub, Brazil's primary production and export region. Hybrids AF-13395 and Goldmine showed the highest m + a′ and d for MMI, indicating suitability for line extraction, as they exhibited high yield and fruit number. These hybrids showed similar m + a′ and non-significant d for soluble solids content (Table 4). Hybrids Gladial and Mandacaru showed the highest m + a′ values for MMI, with non-significant d values.

Deriving m + a′ and d estimates from F1 and F2 means identifies superior hybrids suitable for inbred line development. This methodology identified AF-13395 and Goldmine as optimal hybrids and supported early identification of potential inbred lines as homozygosity increases. Early selection reduces population development costs and enables evaluation of more populations in less time than conventional traditional methods, providing an efficient and cost-effective alternative.

5. Conclusions

In the melon F1 generation, homozygous loci (m + a′) contributed substantially to fruit number, mean fruit mass, yield, flesh thickness, flesh firmness, and soluble solids. Assessments of m + a′ and d quantified the suitability of single-cross hybrid for melon inbred line development. AF-13395 and Goldmine were the most suitable hybrids for superior line selection.

Acknowledgements

This study was partly funded by the Brazilian Coordination for the Improvement of Higher Education Personnel (CAPES, Finance Code 001).

Data Availability Statement

Data are available upon request.

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Edited by

  • Editor:
    Takako Matsumura Tundisi

Publication Dates

  • Publication in this collection
    10 Apr 2026
  • Date of issue
    2026

History

  • Received
    16 Nov 2025
  • Accepted
    02 Feb 2026
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This is an Open Access article distributed under the terms of the Creative Commons Attribution license (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
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