Selection of carioca common bean lines for fusarium wilt resistance, yield, and grain qualityAbstract
The objective of this work was to estimate genetic parameters for the selection of carioca common bean lines resistant to fusarium wilt, with a high yield, an optimal 100-seed weight, and a desirable grain color. Two experiments were conducted during the 2019 and 2020 winter seasons in areas under a high natural pathogen infestation, using a randomized complete block design with three replicates. Sixty lines derived from the cross between cultivars BRS FC402 (resistant) and BRS FC406 (susceptible), plus four checks, were evaluated for fusarium wilt, yield, 100-seed weight, and grain color. Analyses of variance were carried out, and estimates of heritability, expected selection gains, and trait correlations were obtained. Significant differences were observed among lines for all traits. Heritability estimates were 74, 73, 91, and 52% for fusarium wilt, yield, 100-seed weight, and grain color, respectively, indicating a high selection potential. In addition, the reaction to fusarium wilt showed a negative correlation with yield (-0.60) and 100-seed weight (-0.52). Simultaneous selection identified 15 promising lines, generating gains of 13.3, 7.9, 2.4, and 3.5% for fusarium wilt, yield, 100-seed weight, and grain color, respectively. Among these lines, CNFC 20188, CNFC 20134, CNFC 20160, and CNFC 20142 combine all four desirable traits, standing out as promising candidates for the release of new cultivars resistant to fusarium wilt, with a high yield, an optimal 100-seed weight, and a desirable grain color.
Index terms:
Fusarium oxysporum f; sp; phaseoli; Phaseolus vulgaris; heritability; selection gain.
Resumo
O objetivo deste trabalho foi estimar parâmetros genéticos para seleção de linhagens de feijão-carioca resistentes à murcha-de-fusário, com elevada produtividade, massa de 100 grãos ideal e coloração de grãos desejável. Foram conduzidos dois experimentos durante as safras de inverno de 2019 e 2020, em áreas com alta infestação natural do patógeno, tendo-se utilizado delineamento em blocos ao acaso, com três repetições. Sessenta linhagens provenientes do cruzamento entre as cultivares BRS FC402 (resistente) e BRS FC406 (susceptível), além de quatro testemunhas, foram avaliadas quanto à reação à murcha-de-fusário, produtividade, massa de 100 grãos e coloração de grãos. Foram realizadas análises de variância, tendo-se obtido estimativas de herdabilidade, ganhos esperados com a seleção e correlações entre caracteres. Observou-se diferença significativa entre as linhagens para todos os caracteres. As estimativas de herdabilidade foram 74, 73, 91 e 52% para murcha-de-fusário, produtividade, massa de 100 grãos e coloração de grãos, respectivamente, o que indica alto potencial de seleção. Além disso, a reação à murcha-de-fusário apresentou correlação negativa com produtividade (-0,60) e massa de 100 grãos (-0,52). A seleção simultânea identificou 15 linhagens promissoras, com ganhos de 13,3, 7,9, 2,4 e 3,5% para murcha-de-fusário, produtividade, massa de 100 grãos e coloração de grãos, respectivamente. Entre estas linhagens, CNFC 20188, CNFC 20134, CNFC 20160 e CNFC 20142 combinaram os quatro caracteres desejáveis, tendo-se destacado como candidatas promissoras para o lançamento de novas cultivares resistentes à murcha-de-fusário, com elevada produtividade, massa de 100 grãos ideal e coloração de grãos desejável.
Termos para indexação:
Fusarium oxysporum f; sp; phaseoli; Phaseolus vulgaris; herdabilidade; ganho de seleção.
Introduction
Common bean (Phaseolus vulgaris L.) is a legume of great economic, nutritional, and social relevance, being the main source of plant protein for many families and a key source of income for numerous farmers (Izquierdo et al., 2018). However, the production of common bean can be affected by several factors, including inadequate crop management, the use of uncertified and low-quality seeds, unfavorable climatic conditions, and the incidence of diseases (Karavidas et al., 2022).
Fusarium wilt, caused by the fungus Fusarium oxysporum f. sp. phaseoli, stands out among the most important diseases, leading to yield losses of up to 80% (Cavalheiro et al., 2023a). The infection process begins in roots, followed by colonization and obstruction of xylem vessels, resulting in leaf wilting, vascular discoloration, chlorosis, stunting, and premature plant death (Benchimol-Reis et al., 2023).
The use of resistant cultivars is the most efficient control method, as it is not only effective and cost-efficient but also environmentally sustainable (Guimarães & Souza, 2019). The development of these cultivars starts with the identification of resistant plants, followed by the assessment of the nature and magnitude of the genetic effects involved in the observed resistance. In this context, the estimation of genetic parameters is essential to quantify the available variability and to predict the expected gain from selection, guiding breeders in defining optimal selection strategies (Ramalho et al., 2012).
In the literature, several studies have reported the existence of genetic variability for reaction to fusarium wilt (Pereira et al., 2011, 2020; Torres et al., 2021, 2022, 2024; Cavalheiro et al., 2023a, 2023b). Regarding genetic control, some works have indicated that the inheritance of resistance may be oligogenic (Cândida et al., 2009; Batista et al., 2017; Cavalheiro et al., 2023a), while others suggest a polygenic basis (Torres et al., 2022, 2024; Chiwina et al., 2023), reflecting the complexity of the host-pathogen interaction.
For common bean, there are registers of high heritability estimates, ranging from 77 to 94% for resistance to fusarium wilt (Cândida et al., 2009; Pereira et al., 2011; Torres et al., 2022, 2024; Cavalheiro et al., 2023a), as well as of expected selection gains, varying from 23 to 37% (Cândida et al., 2009; Torres et al., 2022, 2024; Cavalheiro et al., 2023a). However, most estimates were calculated under controlled conditions, where resistance expression is less influenced by environmental factors. Therefore, the estimation of genetic parameters under diverse field environments remains limited, restricting the ability of breeding programs to predict selection response and to develop cultivars with a durable resistance. In this scenario, estimating genetic parameters and evaluating resistance under representative conditions are critical steps toward improving selection efficiency.
In the case of carioca-type common bean, in addition to pathogen resistance, cultivars must exhibit market-preferred traits such as a high yield, an optimal 100-seed weight, and a very light-colored beige seed coat with light brown stripes (Cavalheiro et al., 2023a). The greatest challenge for breeding programs lies in developing lines that combine all desirable traits in a single phenotype.
The objective of this work was to estimate genetic parameters for the selection of carioca common bean lines resistant to fusarium wilt, with a high yield, an optimal 100-seed weight, and a desirable grain color.
Materials and Methods
The experiments were conducted at the experimental area of Embrapa Arroz e Feijão, located in the municipality of Santo Antônio de Goiás, in the state of Goiás, Brazil (16º30’17”S, 49º16’53”W, at an altitude 819 m). The local climate is Aw, a tropical savanna in the megathermal zone, according to Köppen, and the soil is classified as a Latossolo Vermelho-Escuro (Santos et al., 2025), corresponding to a positively charged Oxisol (Soil Survey Staff, 2022). The area, used in several previous studies, is characterized by a high natural infestation of Fusarium oxysporum f. sp. phaseoli (Pereira et al., 2018, 2019, 2020; Torres et al., 2021, 2022, 2024; Cavalheiro et al., 2023a, 2023b).
To obtain the breeding lines, a segregating population was derived from the cross between the BRS FC402 and BRS FC406 carioca-type cultivars. Cultivar BRS FC402 has a high resistance to fusarium wilt and anthracnose, a high yield, and a 100-seed weight of 26 g (Melo et al., 2017). Comparatively, cultivar BRS FC406 presents a high yield, resistance to anthracnose and angular leaf spot, and a 100-seed weight of 28 g, but is susceptible to fusarium wilt (Pereira et al., 2021). The population was advanced up to the F4 generation by the bulk method, between 2015 and 2017, in an experimental area naturally infested with the pathogen.
In the F5 generation, during the 2018 winter season, 60 individual plants were randomly harvested from the population. The lines originating from these plants were evaluated in the first field trial in the 2019 winter season (sowing in May), in addition to four carioca-type check cultivars. Seeds harvested from this trial were used for a second experiment, in the same design, in the 2020 winter season (sowing also in May). Cultivars BRS FC402 and BRS Notável were used as resistant checks for fusarium wilt, while BRS Cometa and BRS FC406 were the susceptible checks (Melo et al., 2017; Pereira et al., 2016, 2018, 2021).
The trials were arranged in a randomized complete block design, with plots consisting of three rows, each 3.0 m long, spaced 0.50 m apart. The evaluated traits were: grain yield, 100-seed weight, reaction to fusarium wilt, and grain color after harvest. Grain yield (grams per plot), adjusted to 13% moisture, was obtained by weighing the grains harvested from all plants in the three rows, being later converted to kilogram per hectare. The trait 100-seed weight (grams) was determined by weighing a random sample of 100 seeds from each plot. Reaction to fusarium wilt was assessed 80 days after emergence using the rating scale adapted from Pastor-Corrales & Abawi (1987), which ranged from 1 to 9, representing 0, 0.1-5.0, 5.1-10, 10.1-20, 20.1-40, 40.1-60, 60.1-80, 80.1-90, and 90.1-100% susceptibility in the plot, respectively (Torres et al., 2022). Grain color was evaluated 30 days after harvest using the scale adapted from Silva et al. (2008), which ranged from 1 (very light beige grain coat with light brown stripes) to 5 (dark beige grain coat with dark brown stripes).
The data were subjected to individual and combined analyses of variance, considering only the environment effect as fixed. Prior to both analyses of variance, the assumptions of normality of residuals and homogeneity of variances were verified. Coefficients of variation and selective accuracy were calculated to assess the informativeness of the experiments (Resende & Duarte, 2007). From the analyses of variance, the components of phenotypic variance, genetic variance, genotype × environment interaction variance, and heritability were estimated for all traits, according to the models described by Ramalho et al. (2012). Expected genetic gain was calculated for each trait using a selection intensity of 25%. In addition, the expected gain from the simultaneous selection of traits, at the same intensity, was obtained using the method of independent culling levels (Hazel & Lush, 1942), considering the following selection criteria: fusarium wilt reaction score ≤ 3, grain color score ≤ 3, yield above the mean of 2,089 kg ha-1 of the lines, and 100-seed weight above 29 g. Phenotypic correlations among the evaluated traits were estimated as described by Ramalho et al. (2012), whereas statistical analyses were performed using the GENES software (Cruz, 2013).
Results and Discussion
The estimates of the coefficients of variation obtained in the individual analyses in 2019 and 2020 were: 20 and 18.7% for grain yield, 2.8 and 3.9% for 100-seed weight, and 20.7 and 14.9% for grain color after harvest, respectively (Table 1). These values indicate an adequate experimental precision, consistent with reports in the literature for the same traits (Pereira et al., 2021; Silva et al., 2023). Specifically for reaction to fusarium wilt, the estimates of the coefficients of variation reached higher values of 29.4 and 24.1% in each year, as expected in field evaluations due to a naturally heterogeneous pathogen distribution in the soil (Cavalheiro et al., 2023a, 2023b; Torres et al., 2021, 2022, 2024). In both years, selective accuracy estimates were high (>0.85), showing a good experimental informativeness and strong correlation between genotypic and phenotypic values (Resende & Duarte, 2007).
Summary of individual and combined analyses of variance for grain yield (GY), 100-seed weight (100SW), reaction to fusarium wilt (FOP), and grain color after harvest (GC) of carioca common bean (Phaseolus vulgaris) lines evaluated in 2019 and 2020, in the municipality of Santo Antônio de Goiás, in the state of Goiás, Brazil.
The combined analysis of variance revealed significant differences among lines for all traits, indicating the presence of genetic variability. In the case of reaction to fusarium wilt, the differences among lines confirmed the high disease pressure in the experimental area, consistent with the findings of previous studies conducted at the same site (Pereira et al., 2016, 2018, 2019, 2020; Torres et al., 2021, 2022, 2024; Cavalheiro et al., 2023a, 2023b). These results are important since variability is a prerequisite for breeding, enabling the selection of superior genotypes for fusarium wilt resistance, optimal yield, and ideal grain size and appearance.
The effect of year was significant for all traits, with a considerable variation between means in 2019 and 2020, as follows: 2,516 and 1,636 kg ha-1 for yield, 28.4 and 29.9 g for 100-seed weight, 3.6 and 2.8 for reaction to fusarium wilt, and 2.6 and 3.4 for grain color after harvest, respectively. These results suggest that, although both experiments were carried out at the same site and cropping season, climatic variations across years strongly influenced pathogen behavior and the phenotypic expression of the studied lines.
A significant interaction between lines and years was observed for yield, 100-seed weight, and grain color, which is expected for quantitative traits under a strong environmental influence (Batista et al., 2017). In contrast, reaction to fusarium wilt showed no significant line × year interaction (p>0.05), as reported by Pereira et al. (2019) and Torres et al. (2022), indicating that genotypes classified as superior for resistance remain stable across years and can be reliably selected. This stability may reflect a relatively uniform disease pressure in the experimental area, possibly due to the predominance of a single or limited number of Fusarium oxysporum f. sp. phaseoli races. This information is not yet known since there are still gaps regarding the distribution of the pathogen (Benchimol-Reis et al., 2023), specifically of its physiological races reported worldwide, confirming its wide genetic variability (Henrique et al., 2015). Furthermore, the consistent response observed may suggest resistance controlled by major-effect loci, as proposed by Batista et al. (2017).
In both individual and combined analyses, genetic variance accounted for a large proportion of phenotypic variance across all evaluated traits (Table 2), indicating that most of the observed variation was predominantly genetic in origin. In the joint analysis, genetic variance consistently exceeded the genotype × environment interaction variance for all traits, suggesting that the performance of the lines was only slightly influenced by environmental variation across years. Similar results have been obtained for common bean, especially regarding reaction to fusarium wilt, indicating a consistent resistance expression across environments, while highlighting genetic variance as the main driver of phenotypic differences among lines (Torres et al., 2022; Cavalheiro et al., 2023a). Estimates of phenotypic, genetic, and environmental variance are fundamental for breeding programs, as they allow quantifying the relative contribution of genetic and environmental factors to trait expression, guiding selection strategies (Ramalho et al., 2012).
Estimates of the genetic parameters and simultaneous selection gain, at a 25% selection intensity, for grain yield (GY), 100-seed weight (100SW), reaction to fusarium wilt (FOP), and grain color after harvest (GC) of carioca common bean (Phaseolus vulgaris) lines in 2019 and 2020.
For reaction to fusarium wilt, the heritability estimates in each environment were lower than those of the combined analysis, indicating a more reliable selection when data from both years are considered jointly (Table 2). Cavalheiro et al. (2023a) also reported a high heritability of 76.6% for resistance to fusarium wilt in carioca bean. Torres et al. (2022) obtained even higher estimates of 90.9% for black bean genotypes, which tend to show higher resistance levels than carioca genotypes according to Pereira et al. (2016).
Grain yield showed a high heritability in the combined analysis (Table 2), which is particularly notable since this trait is quantitatively inherited and strongly influenced by environmental fluctuations. The variation observed between years reinforces the importance of multienvironment evaluations to improve selection accuracy for yield in common bean breeding programs (Ramalho et al., 2012). The trait 100-seed weight also presented a consistently high heritability, indicating a strong genotype-phenotype association and a greater predictability of selection response. For grain color after harvest, the estimates were moderate as in previous reports, reflecting the combined influence of genetic control and environmental effects during grain maturation and storage (Cavalheiro et al., 2023a). These results are especially relevant because they were obtained under field conditions, confirming that genetic variance remains sufficiently high to support a consistent selection across years.
Due to the expected gain from direct selection, the 15 selected lines, at a 25% selection intensity, reduced the mean score for reaction to fusarium wilt by 0.58 in the joint analysis, which corresponds to an 18% gain (Table 2). The obtained value was lower than those reported in the literature of 26.8 to 35.5% under controlled conditions (Cândida et al., 2009), likely due to the greater environmental variability in the experimental field, which provides a more realistic estimate of selection response for breeding programs. These findings are important because selection gain represents the targeted shift in the mean of a trait toward desirable levels, maximizing the frequency of favorable alleles that control it (Ramalho et al., 2012).
Among the 60 lines evaluated, 40 had mean scores below 3 (Table 3), being classified as resistant (Pastor-Corrales & Abawi, 1987). This probably occurred because the population was advanced using the bulk method during five generations in a field naturally infested with the pathogen, which may have facilitated the elimination of the most susceptible plants in the early segregating generations. This result reflects the high standard of the studied population, which reduces the likelihood of highly expressive gains from selection, given the high frequency of favorable alleles conferring resistance.
Means based on the joint analysis of 60 common bean lines (Phaseolus vulgaris) and 4 checks for grain yield (GY), 100-seed weight (100SW), reaction to fusarium wilt (FOP), and grain color after harvest (GC)(1).
The direct selection gains obtained for grain yield, 100-seed weight, and grain color in the joint analysis are highly relevant for breeding programs (Table 2). This is meaningful considering that yield and grain-quality traits have a complex inheritance, being controlled by multiple genes subject to epistatic interactions, pleiotropic effects, multiple allelism, and linkage, which hinder their selection (Ramalho et al., 2012). The aforementioned result is an indicative that the obtained gains represent a greater alignment of the selected lines with market requirements, favoring product commercialization.
The gains obtained through simultaneous selection, although lower than those resulting from direct selection, were satisfactory for all traits in the joint analysis (Table 2). This favored an increase in the frequency of favorable alleles for multiple traits, which will reduce the expended time and costs of breeding programs. Similar results were reported by Cavalheiro et al. (2023a), who observed gains of 12.9% for resistance to fusarium wilt, 5.6% for grain yield, 3.4% for 100-seed weight, and 6.1% for grain color. Torres et al. (2022) found higher gains of 22.1 and 7.7% for resistance to fusarium wilt and 100-seed weight, respectively, and similar results of 6.6% for yield.
For reaction to fusarium wilt, all 60 lines were superior to the two susceptible check cultivars, BRS Cometa and BRS FC406 (Table 3), confirming the success of natural selection during generational advancement. In relation to the BRS Notável and BRS FC402 resistant checks, 58 and 39 lines, respectively, achieved equal or lower scores.
The BRS FC402 parental cultivar carries the same quantitative trait loci (QTL) associated with resistance to fusarium wilt identified in BRS FP403 (Torres et al., 2024; Ciappina et al., 2026), a black bean cultivar recognized for its resistance to the disease. This QTL, designated FOP2.3403H and located on chromosome 2, explained 40.5% of phenotypic variance (Torres et al., 2024). The studied cultivar, however, does not carry the QTLs associated with resistance in other sources, including: FOP7.1NS, on chromosome 7 in the BRS Notável resistant cultivar check, explaining 45 of phenotypic variance (Cavalheiro, 2021); and FOP8.1EH, on chromosome 8 in the BRS Esplendor black bean cultivar (Ciappina et al., 2026), explaining 66%. These results underscore the value of the selected lines as promising material for resistance-allele pyramiding and for the development of common bean cultivars with a stable resistance to fusarium wilt.
Regarding yield traits, 14 lines surpassed the highest-yielding check cultivar, BRS FC402, showing a high yield potential (Table 3). For 100-seed weight, the minimum value for the selected lines was 29.1 g, exceeding that of all checks and meeting the commercial standards of a grain weight above 25 g (Pereira et al., 2012).
For grain color after harvest, the selected line with the highest score of 3.0 produced lighter grains than three checks (cultivars BRS Notável, BRS Cometa, and BRS 406), indicating conformity with commercial standards (Table 3). However, the grains produced by these cultivars presented a light beige background and dark brown stripes, while the market has increasingly preferred grains with a very light beige background but light brown stripes (Cavalheiro et al., 2023a). Considering these standards, lines CNFC 20188, CNFC 20149, and CNFC 20175 stood out for color selection, with a score equal to or lower than that of the BRS FC402 resistant check cultivar, which has light beige grains with light brown stripes, in line with market values and industry preferences (Melo et al., 2017).
Overall, line CNFC 20188 stood out from the others, surpassing all checks for the four traits evaluated. Lines CNFC 20134, CNFC 20160, and CNFC 20142 also showed a high potential, outperforming cultivar BRS FC402 for fusarium wilt resistance, yield, and 100-seed weight, with a 2.7 score for grain color. In the future, it is essential to evaluate these lines across multiple environments to validate their potential for release as commercial cultivars.
The correlation estimates indicated that the scores for reaction to fusarium wilt were associated with a yield of -0.60 and 100-seed weight of -0.52. These associations were negative and significant (p<0.01), showing that plants with a higher disease incidence, i.e., higher scores, exhibited lower yields. Conversely, plants with a better health contributed to higher yields and a consequent heavier seed weight. Cavalheiro et al. (2023a) and Torres et al. (2022) also reported negative correlations of -0.58 and -0.72 between yield and reaction to fusarium wilt, respectively, which are close to the values obtained in the present study. For 100-seed weight and reaction to fusarium wilt, other authors found estimates of -0.36 (Cavalheiro et al., 2023a) and -0.28 (Torres et al., 2022), both lower than those presented here.
Conclusions
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1. Carioca common bean (Phaseolus vulgaris) lines present genetic variability among themselves, as well as high heritability estimates and expected selection gains for reaction to fusarium wilt, grain yield, 100-seed weight, and grain color after harvest.
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2. Fifteen lines combine resistance to fusarium wilt, high yield, heavier 100-seed weight, and grain color meeting market requirements, among which CNFC 20188, CNFC 20134, CNFC 20160, and CNFC 20142 are the most outstanding.
Acknowledgments
To Embrapa Arroz e Feijao and its partners for providing infrastructure and labor support; to Coordenacao de Aperfeicoamento de Pessoal de Nivel Superior (CAPES), for granting a doctoral scholarship to AHC (grant no. 88882.386276/2019-01), and to Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq), for granting a technological and industrial development scholarship to NVC (grant no. 384993/2025-7), as well as for granting scholarships for technological development and innovative extension to HSP and LCM authors.
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Declaration of use of AI technologies
No generative artificial intelligence (AI) was used in this study.
Data availability statement
Data available upon request: research data are only available upon reasonable request to the corresponding author.
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Edited by
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Chief editor:
Edemar Corazza
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Edited by:
Mírian Baptista
