Genetic Variability, Broad-Sense Heritability, and Selection of Superior Genotypes for Fruit Improvement in Platonia insignis
by
, , , , and
Suzane Sá Matos Ribeiro
1, 
Sérgio Heitor Sousa Felipe
1
,
Givago Lopes Alves
1,
Priscila Marlys Sá Rivas
1
,
Juliane Maciel Henschel
2,
Lúcio Rafael Rocha de Moraes
3,
Luís Carlos Ferreira Reis
3,
José Ribamar Gusmão Araújo
1,
Marcos Vinícius Marques Pinheiro
4,
Diego Silva Batista
1,2,*
and
Thais Roseli Corrêa
1,* 1
Programa de Pós-Graduação em Ciências Agrárias, Universidade Estadual do Maranhão, São Luís 65055-310, MA, Brazil
2
Programa de Pós-Graduação em Agronomia, Universidade Federal da Paraíba, Areia 58397-000, PB, Brazil
3
Departamento de Fitotecnia e Fitosanidade, Centro de Ciências Agrárias, Universidade Estadual do Maranhão, São Luís 65055-310, MA, Brazil
4
Campus Frederico Westphalen (URI-FW), Universidade Regional Integrada do Alto Uruguai e das Missões, Frederico Westphalen 98400-000, RS, Brazil
*
Authors to whom correspondence should be addressed.
Int. J. Plant Biol. 2025, 16(3), 108; https://doi.org/10.3390/ijpb16030108
Submission received: 24 July 2025
/
Revised: 9 September 2025
/
Accepted: 10 September 2025
/
Published: 15 September 2025
(This article belongs to the Section Plant Biochemistry and Genetics)
Abstract
Platonia insignis Mart. is a native Amazonian fruit tree with considerable agro-industrial potential, yet it remains underutilized due to limited domestication efforts and the absence of breeding programs or improved genetic lines. This study aimed to estimate genetic parameters based on morpho-agronomic fruit traits and to identify superior genotypes from natural coastal populations in the Brazilian Amazon. Thirteen genotypes were evaluated for 16 biometric and compositional traits. Genetic parameters were estimated using REML/BLUP (Restricted Maximum Likelihood/Best Linear Unbiased Prediction) procedures, and a rank–sum selection index was applied to identify elite individuals. The results revealed substantial phenotypic and genetic variability, with broad-sense heritability values ranging from moderate to high for key traits, including longitudinal fruit diameter (0.81), fruit fresh mass (0.66), and seed fresh mass (0.75). Selection accuracy was high (≥0.96) across most traits, indicating strong experimental reliability. Genotypic correlations highlighted favorable associations among traits related to pulp yield, sugar content, and seed reduction. Six superior genotypes (G7, G1, G6, G3, G2, and G4) exhibited optimal profiles for fruit quality and productivity. These findings provide a strong foundation for breeding strategies and support the genetic conservation and domestication of P. insignis as a native species of high economic and ecological importance.
1. Introduction
The Brazilian Amazon biome harbors a remarkable diversity of native fruit tree species with substantial potential for integration into conservation and genetic improvement programs. Initiatives focused on sustainable use, domestication, and genetic enhancement of these species can significantly contribute to biodiversity conservation, food security, and public health, as many fruits are rich in fiber, vitamins, and nutraceutical compounds [1,2]. However, species such as Platonia insignis Mart. (Clusiaceae), which hold high ecological and socio-economic value, are increasingly threatened by predatory extraction, agricultural expansion, and environmental degradation, resulting in the erosion of genetic resources and the loss of adaptive potential [3,4].
P. insignis is a multipurpose species valued for both its timber and fruit. Its pulp is widely consumed fresh or processed into sweets, juices, jams, and ice cream, and is recognized for its high market value and functional properties [5,6]. Despite this economic relevance, the fruit production chain remains underdeveloped and largely dependent on extractivism, with limited efforts directed toward the development of genetically improved cultivars for commercial cultivation [3]. Recent studies have revealed pronounced genetic structuring and low diversity in Amazonian populations of P. insignis, underscoring the urgency of conservation strategies and the selection of adapted genotypes, particularly across contrasting environments such as the Amazon and Cerrado biomes [4]. Moreover, recent advances have expanded knowledge of the biology, ecology, and potential uses of the species while also identifying critical gaps for genetic improvement and domestication [6].
Although Platonia insignis has high economic value, its cultivation remains incipient, with fruit production primarily dependent on extractivism from natural populations. In Pará state, Brazil, early cultivation initiatives have been reported, including the management of natural regrowth and the planting of seedlings derived from both seeds and grafted plants [7]. Grafted seedlings reduce the juvenile period to four to six years, whereas seed-derived plants begin fruiting only after ten to thirteen years [5]. However, commercial adoption is still limited by the lack of efficient propagation techniques, the species’ dependence on cross-pollination, and restricted access to elite germplasm [5,6]. As a result, exploitation continues to rely heavily on wild populations, raising concerns about the long-term sustainability of its genetic resources.
In this context, the estimation of genetic parameters and the phenotypic characterization of fruit traits in wild populations are crucial steps toward the domestication of native species. These approaches enable the identification of individuals with high agronomic potential and support the development of elite cultivars for cultivation and breeding purposes [8,9]. Studies on other underutilized species, such as Cordia myxa, have demonstrated that multivariate analyses of morphological traits can uncover significant diversity and guide early selection [10]. Similarly, in Endopleura uchi, moderate heritability values and a strong negative correlation between pulp yield and fruit diameter indicate both genetic variability and selection potential [11].
The selection of accessions based on fruit quality and pulp yield has also proven effective in economically important native species such as Euterpe edulis. Recent studies have reported high genetic variability and substantial predicted gains for key traits, including total anthocyanin content and pulp percentage, thereby supporting the selection of superior genotypes for commercial cultivation and breeding programs [12]. These findings highlight the relevance of morpho-agronomic descriptors and robust statistical approaches, such as REML/BLUP and rank-based selection indices, in identifying elite trees under early selection schemes.
Given this background, the present study aimed to assess genetic variability and broad-sense heritability of fruit morpho-agronomic traits and to identify superior genotypes of P. insignis for fruit improvement. The evaluation was based on wild individuals from natural coastal populations in the Brazilian Amazon, with the goal of supporting conservation initiatives and advancing the domestication of this native species.
2. Results
2.1. Genetic Parameters Based on Morpho-Agronomic Traits of Platonia insignis Fruits
The estimation of genetic parameters revealed considerable variation in the relative contributions of genetic and environmental effects across the evaluated traits. Genotypic variance (Vg) exceeded environmental variance (Ve) for five traits—longitudinal fruit diameter (LFD), pericarp thickness (FST), LFD/TFD ratio, fruit fresh mass (FFM), and seed fresh mass (SFM)—indicating that their phenotypic expression is predominantly controlled by genetic factors and, therefore, more amenable to selection. In contrast, traits such as transversal fruit diameter (TFD), fruit color (FC), peel fresh mass (PFM), pulp fresh mass (PuFM), number of seeds per fruit (NSF), number of parthenocarpic segments per fruit (NPSF), peel yield (PY), pulp yield (PUY), seed yield (SY), soluble solids content (SSC), and titratable acidity (TTA) exhibited Ve values greater than Vg, suggesting stronger environmental influence on their variation (Table 1). Overall, environmental effects exerted a greater influence on most traits, as evidenced by the predominance of Ve over Vg in 11 of the 16 parameters evaluated.
Broad-sense heritability (h2g) values were moderate to high for several traits, with particularly high estimates for LFD (0.81), FFM (0.66), and SFM (0.75), supporting their potential utility in fruit improvement strategies. Moderate heritability was also observed for FST (0.57), LFD/TFD ratio (0.54), and PuFM (0.40), indicating consistent genetic control. In contrast, traits such as pulp yield (PUY), seed yield (SY), titratable acidity (TTA), and SSC exhibited low or negligible heritability (h2g < 0.10), suggesting that their expression is more strongly influenced by environmental factors.
Selective accuracy (Acc) values were uniformly high for most traits (≥0.96), indicating strong experimental precision and reliable prediction of genetic values. These results reinforce the robustness of the phenotypic assessments and support the reproducibility of selection outcomes across breeding cycles.
The coefficients of relative variation (CVr) confirmed substantial genetic variability among genotypes. High CVr values were observed for LFD (27.47), FFM (27.70), FST (24.02), and SFM (21.02), indicating strong potential for genetic gain, particularly in the early stages of domestication and breeding programs. In contrast, low CVr values for SY (0.11), TTA (0.07), and PUY (0.66)—combined with their low broad-sense heritability and selection accuracy estimates—suggest that these traits are predominantly influenced by environmental factors under the evaluated conditions, thereby limiting their utility in early selection strategies.
Among these, soluble solids content (SSC) exhibited one of the lowest genotypic variance estimates (Vg = 1.00), low broad-sense heritability (h2g = 0.10), and only moderate selection accuracy (Acc = 0.67), indicating strong environmental influence on this trait. These results point to limited potential for improving SSC through selection under the current experimental conditions. Overall, environmental effects played a more prominent role in shaping phenotypic variability, as reflected by the predominance of Ve over Vg in most of the traits evaluated (11 of 16).
Detailed means and standard deviations for each morpho-agronomic trait across the 13 genotypes are provided in Supplementary Table S1. The significance of genotypic variance (Vg) was assessed using likelihood ratio tests (LRTs), with results summarized in Table 2.
2.2. Phenotypic Traits of Shape and Color in Platonia insignis Fruits
The study revealed substantial genetic variability among the genotypes, as reflected by the diverse phenotypic patterns in fruit morphology (Figure 1). Notably, variation in fruit color and shape showed no clear association with the geographic origin of the genotypes.
In terms of external color, genotype G7 exhibited a red–orange hue, whereas G5, G8, and G10 were predominantly yellow. Genotypes G2 and G3 displayed a yellow–orange color; G12 showed a yellow–orange–green tone; G1, G4, and G6 were orange; G9 and G13 presented orange–yellow coloration; and G11 exhibited a green–yellow appearance (Figure 1).
With respect to fruit shape, a predominance of flattened forms was observed in genotypes G1, G4, G6, G8, G10, G12, and G13. In contrast, G2, G3, G5, G7, G9, and G11 exhibited rounded fruits (Figure 1). Although the fruit apex was not quantitatively assessed, some genotypes—such as G1, G6, G7, and G12 (Figure 1A, Figure 1F, Figure 1G, and Figure 1M, respectively)—displayed a distinctly acuminate apex, visually distinguishing them from the others.
2.3. Genotypic Correlations Based on Morpho-Agronomic Traits of Platonia insignis Fruits
The genotypic correlation analysis revealed significant associations among the biometric, compositional, and quality-related traits of P. insignis fruits (Figure 2). These correlations highlight underlying genetic interdependence and provide valuable insights for guiding indirect selection strategies in early breeding programs.
A very strong positive correlation was observed between fruit fresh mass (FFM) and pericarp thickness (FST) (r = 0.90, p < 0.01), indicating that heavier fruits tend to have thicker structural tissues. In contrast, a very strong negative correlation was detected between seed fresh mass (SFM) and number of seeds per fruit (NSF) (r = −0.96, p < 0.01), suggesting that fruits with more seeds tend to produce smaller or lighter individual seeds, reflecting a trade-off in seed biomass allocation.
Several other strong positive correlations further reinforced key relationships. FFM was positively correlated with the number of parthenocarpic segments per fruit (NPSF) (r = 0.76, p < 0.01), suggesting that fruit mass increases with the degree of segmentation. Pulp fresh mass (PuFM) showed strong and significant positive associations with NSF (r = 0.86, p < 0.01), longitudinal fruit diameter (LFD) (r = 0.77, p < 0.01), and soluble solids content (SSC) (r = 0.73, p < 0.01). Additionally, NSF was strongly positively correlated with LFD (r = 0.83, p < 0.01), reinforcing the link between seed abundance and fruit size, while SSC was positively associated with titratable acidity (TTA) (r = 0.74, p < 0.01).
Strong negative correlations were also observed between PuFM and SFM (r = −0.79, p < 0.01), and between SFM and LFD (r = −0.87, p < 0.01). These results indicate that greater pulp accumulation is genetically associated with reduced seed mass, and that fruits with larger locular diameters tend to bear seeds with lower total biomass.
2.4. Genotypes Selection by Rank-Sum Index
The rank–sum index revealed significant differences in performance among the top six genotypes of P. insignis (Table 3). Genotype G7 exhibited the highest overall performance, with the lowest average rank (4.92), followed by G1, G6, G3, G2, and G4. These genotypes consistently achieved superior rankings for traits prioritized in pulp yield and fruit quality improvement.
Genotype G7 ranked first for pulp fresh mass (PuFM), number of seeds per fruit (NSF), and number of parthenocarpic segments per fruit (NPSF), and second for soluble solids content (SSC), indicating a favorable combination of high pulp accumulation, reduced seed content, and elevated sugar levels.
Genotype G1 also performed strongly, securing second place in seed fresh mass (SFM), PuFM, and pulp yield (PUY), highlighting its potential for pulp productivity. G6 followed closely, with an average rank of 6.17, showing consistent performance across traits, including fourth in PuFM and fifth in NSF, reflecting a balanced profile for fruit composition and yield.
Genotypes G3 and G2 had average ranks of 6.25 and 6.33, respectively. G3 performed well for peel fresh mass (PFM) and peel yield (PY), although it ranked lower for PUY and SSC. In contrast, G2 showed top rankings for PFM (1st), SFM (1st), and pericarp thickness (FST) (2nd), but weaker performance in PUY (10th) and SSC (13th), suggesting a structure-oriented profile with limited sweetness and pulp proportion.
Genotype G4, with an average rank of 6.50, completed the top group, exhibiting strong performance for titratable acidity (TTA) (1st) and seed yield (SY) (2nd), but modest rankings for most pulp-related traits.
2.5. Principal Component Analysis of Morpho-Agronomic Traits
To explore the overall structure of phenotypic variation among P. insignis genotypes, a principal component analysis (PCA) was performed using 16 morpho-agronomic fruit traits. The first two principal components (PC1 and PC2) explained 39.9% and 21.4% of the total variance, respectively (Figure 3A). Together, they captured 61.3% of the observed variability, providing a reliable multivariate representation of trait dispersion.
The biplot analysis revealed two distinct genotype groups, clearly separated along the PC1 axis. Group 1 (red) comprised genotypes with higher seed fresh mass (SFM) and moderate peel yield (PY), whereas Group 2 (blue) included genotypes with superior pulp fresh mass (PuFM), longitudinal fruit diameter (LFD), and fruit fresh mass (FFM), as well as favorable values for soluble solids content (SSC) and number of parthenocarpic segments per fruit (NPSF).
The contributions of individual traits to PC1 indicated that PuFM, LFD, FFM, SFM, and number of seeds per fruit (NSF) were the main discriminating descriptors (Figure 3B), highlighting their importance in explaining intergenotypic variation. In contrast, PC2 was primarily influenced by peel fresh mass (PFM), fruit color (FC), and peel yield (PY) (Figure 3C), suggesting that these traits contribute to secondary differentiation and may reflect environmental or physiological effects not fully captured by PC1.
3. Discussion
Accessing genetic parameters in natural populations of a species is a critical initial step for identifying promising genotypes for both genetic conservation and plant breeding programs [9,13]. Our results demonstrated favorable genetic estimates for several morpho-agronomic fruit traits in P. insignis, reinforcing the feasibility of initiating a breeding program for this native species. Notably, this is the first study to report the selection of six elite genotypes (G7, G1, G6, G3, G2, and G4) with potential for improving fruit quality and pulp production in coastal populations of the Brazilian Amazon.
The significant genetic and phenotypic variability observed is consistent with expectations for a species with open pollination and sporophytic self-incompatibility, particularly in populations located within the Amazonian center of diversity [5]. Such reproductive and geographic factors contribute to high heterozygosity and variability in natural populations. Additionally, the influence of environmental heterogeneity on trait expression must be considered, as highlighted by Leão et al. [14].
Moderate to high broad-sense heritability estimates for key morpho-agronomic traits confirm that genetic factors play a substantial role in trait variation, supporting their use in selection strategies. Similar findings were reported by Santos et al. [15] in Psidium cattleyanum, which also exhibited high phenotypic variability and elevated heritability for fruit traits. This agreement reinforces the role of outcrossing reproductive systems and natural genetic diversity in shaping tropical populations with domestication potential.
Although some quality traits exhibited low heritability, the overall selection accuracy was high, increasing confidence in early selection, particularly for perennial species with long reproductive cycles [16]. Evangelista et al. [17] also reported high selection accuracy for grain yield in Jatropha curcas using both frequentist and Bayesian approaches, despite low to moderate heritability values. These findings highlight the relevance of combining morpho-agronomic evaluations with robust statistical models for effective early selection in perennial crops.
Trait correlations revealed consistent patterns that support indirect selection strategies. Strong positive associations between pulp fresh mass (PuFM) and traits such as soluble solids content (SSC) and longitudinal fruit diameter (LFD), along with a strong negative correlation with seed fresh mass (SFM), suggest that genotypes combining high pulp accumulation and sweetness with lower seed biomass are particularly desirable. Similar correlations have been reported in Psidium cattleyanum, where strong associations were observed between fruit and pulp mass, and negative correlations occurred between seed number and pulp yield [15]. These results are consistent with the genetic relationships identified in P. insignis, highlighting the effectiveness of using fruit structural and compositional traits to guide the selection of superior genotypes.
Costa et al. [11] employed the rank–sum method to select Endopleura uchi genotypes with superior performance in ten fruit traits. In the present study, genotypes G7, G1, G6, G3, G2, and G4 stood out for their balanced profiles, particularly for traits related to pulp yield and fruit quality, reinforcing the efficiency of this method in identifying individuals with high potential for pulp-focused breeding. However, it is important to note that genotypes not included among the top six should not be disregarded, as they may harbor favorable characteristics for other breeding objectives, such as seed yield, peel traits, or adaptability to specific environments. The reliability of the selection was further enhanced by incorporating genetic accuracy values, strengthening the robustness of the decisions made.
In summary, this research demonstrates the genetic potential of P. insignis and supports its inclusion in breeding and conservation programs. One limitation of this study is the absence of molecular data and environmental control, which may have influenced phenotypic expression and affected the precision of genetic estimates. Future studies should incorporate molecular markers and conduct evaluations under controlled environmental conditions to refine the understanding of genetic variability and improve selection accuracy.
4. Materials and Methods
4.1. Study Area, Plant Material, and Fruit Harvesting
Thirteen non-clonal fruit trees of Platonia insignis were selected from natural populations along the Brazilian Amazonian coast, Maranhão State, Brazil (Figure 4, Table 4). Trees with a diameter at breast height (DBH) ≥ 25 cm during the fruiting period were chosen to ensure reproductive maturity and adequate fruit availability. The sampling areas were selected based on their observed phenotypic fruit diversity (Figure 1). All selected trees were at least 30 m apart to minimize the likelihood of clonality, ensuring that each sampled tree represented an independent genotype.
Fruit harvesting was conducted between December and March 2022, coinciding with the natural fruiting and dispersal period. Twenty-five fruits were collected from each mother-tree genotype (totaling 350 fruits), packed in bags, and transported to the Laboratory of Phytotechnics and Post-Harvest (LAPPH) at the Universidade Estadual do Maranhão, São Luís, MA, Brazil. In this experimental design, each tree was considered an experimental unit, while the fruits collected from the same tree served as replication units. This approach reflects the natural population structure of P. insignis, as no clonal orchards or commercial plantations are yet available for this species.
4.2. Determination of Morpho-Agronomic Traits
The morpho-agronomic characterization of the 25 fruits from each genotype was performed using a digital caliper (Zaas Precision®, Amatools, Piracicaba, SP, Brazil). Measurements included longitudinal fruit diameter (LFD, mm), transversal fruit diameter (TFD, mm)—defined as the distance between peduncle scars and stigma—and pericarp thickness (FST, mm), which comprises the epicarp and mesocarp of the fruit. Additionally, fruits were assigned color scores (FC) ranging from 1 to 9: red–orange (1), yellow (2), yellow–orange (3), yellow–orange–green (4), orange (5), orange–yellow (6), green (7), green–yellow (8), and green–orange (9). Fruit shape was classified as rounded, flat, or oval based on the LFD/TFD ratio, with values from 0.95 to 1.05 [18].
After fruit processing, the fresh mass of the fruit (FFM, g), peel (PFM, g), pulp (PuFM, g), and seed (SFM, g) were measured using a precision electronic scale (Shimadzu®, AUY220, Tokyo, Japan). The number of parthenocarpic segments per fruit (NPSF, unit) and seeds per fruit (NSF, unit) were counted. Parthenocarpic segments were defined as visible internal fruit compartments that did not contain developed seeds, as determined after pulp removal. Peel yield (PY, %), pulp yield (PUY, %), and seed yield (SY, %) were calculated following Guimarães et al. [18], with PUY representing the edible portion of the fruit.
4.3. Determination of Soluble Solids and Titratable Acidity
The processed pulp of Platonia insignis fruits was used for chemical analyses. Soluble solids content (SSC) was determined using an analog refractometer (Model JuanJuan) and expressed in °Brix. For diluted samples, final values were corrected using a dilution factor calculated as the ratio between the total mass (sample + solvent) and the sample mass. Titratable acidity (TTA) was measured by titration with 0.1 M sodium hydroxide solution, using phenolphthalein as an indicator, and results were expressed as a percentage of citric acid equivalent (% citric acid), following the AOAC [19] protocol.
4.4. Determination of Genetic Parameters
Genetic parameters were estimated using Restricted Maximum Likelihood (REML) and Best Linear Unbiased Prediction (BLUP) procedures, based on mixed linear models implemented in the SELEGEN-REML/BLUP software [20]. The analysis employed Model 2, which assumes complete blocks with replications treated as fixed effects.
The statistical model used was
where
y = Xr + Zg + Wp + e
y is the vector of observed data;
r is the vector of replication effects (assumed as fixed) included with the overall mean;
g is the vector of genotypic effects (assumed as random);
p is the vector of plot effects (random);
e is the vector of residual errors (random);
X, Z, and W are the respective incidence matrices for these effects.
The fit of the model and the significance of random effects were evaluated using deviance analysis based on the Likelihood Ratio Test (LRT). From this model, estimates of genotypic variance (Vg), environmental variance (Ve), phenotypic variance (Vf), broad-sense heritability (h2g), heritability on a mean basis (h2mc), selection accuracy (Acc), and the coefficient of relative variation (CVr) were obtained for each evaluated trait [21]. The magnitudes of broad-sense heritability and selection accuracy were classified according to the criteria proposed by Resende et al. [21], as summarized in Table 5.
Deviance analysis was used to compare model fit and assess the significance of random effects, as implemented in the SELEGEN software [22].
4.5. Statistical Analysis
All genetic parameters were estimated using SELEGEN-REML/BLUP (Statistical System and Computerized Genetic Selection by Mixed Linear Models), version 2016, Model 2 [22].
4.6. Multivariate Analyses and Graphs
Pearson’s genetic correlations were calculated using the genotypic values estimated by the SELEGEN-REML/BLUP model (μ + g) and analyzed between variables. Principal component analysis (PCA) was also performed using these genotypic values to explore relationships among traits and identify the variables contributing most to phenotypic divergence. Both analyses were conducted in R Studio (version 2024.12.1+563) [23].
4.7. Selection Index
The rank–sum index was used to classify genotypes according to favorable values for each trait of interest. After this classification, the positions of all traits for each genotype were summed, forming the index originally proposed by Mulamba and Mock [24]. This index was then used to calculate the final selection index and rank the genotypes, following the procedures described by Cruz et al. [25].
5. Conclusions
This study confirms substantial genetic variability and favorable morpho-agronomic performance among Platonia insignis genotypes sampled from natural populations along the Brazilian Amazonian coast. Traits related to fruit size, pulp content, and sugar concentration exhibited moderate to high broad-sense heritability and consistently high selection accuracy, indicating strong genetic control and reliable predictability across selection cycles. The rank–sum index identified six superior genotypes (G7, G1, G6, G3, G2, and G4) that combine a balanced profile of desirable attributes, including high pulp yield, reduced seed mass, and elevated soluble solids. These genotypes are suitable for inclusion in breeding programs targeting both agronomic performance and fruit quality.
These results provide a solid foundation for the development of domestication strategies, the implementation of in situ and ex situ conservation initiatives, and the selection of improved cultivars adapted to regional production systems. Collectively, this work contributes to the genetic valorization of P. insignis, a native Amazonian fruit tree of high ecological and economic importance.
Supplementary Materials
The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/ijpb16030108/s1, Table S1: Mean ± standard deviation of 16 morpho-agronomic and compositional fruit traits across 13 Platonia insignis genotypes from natural Amazonian populations. Data are based on 25 fruits per genotype (n = 25).
Author Contributions
Conceptualization, T.R.C.; methodology, S.S.M.R., T.R.C. and S.H.S.F.; formal analysis, S.S.M.R., L.R.R.d.M., P.M.S.R. and L.C.F.R.; investigation, S.S.M.R., L.R.R.d.M. and L.C.F.R.; resources, T.R.C. and J.R.G.A.; writing—original draft preparation, S.S.M.R., T.R.C., G.L.A. and S.H.S.F.; writing—review and editing, G.L.A., P.M.S.R., J.M.H., D.S.B., M.V.M.P., S.H.S.F. and T.R.C.; visualization, G.L.A., P.M.S.R., J.M.H., D.S.B., S.H.S.F. and T.R.C.; supervision, T.R.C. and S.H.S.F.; project administration, T.R.C. and J.R.G.A.; funding acquisition, T.R.C. and J.R.G.A. All authors have read and agreed to the published version of the manuscript.
Funding
This research was funded by the Fundação de Amparo à Pesquisa e ao Desenvolvimento Científico e Tecnológico do Maranhão—FAPEMA (Foundation for the Support of Research and Scientific and Technological Development of Maranhão), grant number 00675/19.
Data Availability Statement
The original contributions presented in the study are included in the article, further inquiries can be directed at the corresponding author.
Acknowledgments
The authors thank the National Council for Scientific and Technological Development (CNPq, Brasília, DF, Brazil: Grants no. PQ 304214/2022-1 and PQ 307349/2023-3 to Diego S. Batista and Thais R. Corrêa, respectively); the Coordination for the Improvement of Higher Education Personnel (CAPES, Brasília, DF, Brazil: Finance code 001 to Suzane S. M. Ribeiro and the National Postdoctoral Program to Priscila M. S. Rivas).
Conflicts of Interest
The authors declare no conflicts of interest.
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Figure 1.
Phenotypic aspects of fruits from the 13 Platonia insignis genotypes collected from natural populations along the Brazilian Amazonian coast. Each letter corresponds to a specific genotype: (A) G1; (B) G2; (C) G3; (D) G4; (E) G5; (F) G6; (G) G7; (H) G8; (I) G9; (J) G10; (K) G11; (L) G12; (M) G13. Bars: 2 cm.
Figure 1.
Phenotypic aspects of fruits from the 13 Platonia insignis genotypes collected from natural populations along the Brazilian Amazonian coast. Each letter corresponds to a specific genotype: (A) G1; (B) G2; (C) G3; (D) G4; (E) G5; (F) G6; (G) G7; (H) G8; (I) G9; (J) G10; (K) G11; (L) G12; (M) G13. Bars: 2 cm.
Figure 2.
Genotypic correlation coefficients for fruit traits of the 13 Platonia insignis genotypes collected from natural populations on the Brazilian Amazonian coast. Longitudinal fruit diameter (LFD), transversal fruit diameter (TFD), fruits pericarp thickness (FST), fruit color (FC), LFD/TFD ratio, fruit fresh mass (FFM), peel fresh mass (PFM), pulp fresh mass (PuFM), seed fresh mass (SFM), total number of parthenocarpy segments per fruit (NPSF), total number of seeds per fruit (NSF), peel yield (PY), pulp yield (PUY), seed yield (SY), soluble solids content (SSC), and titratable acidity (TTA). * p ≤ 0.05; ** p ≤ 0.01; and ns non-significant.
Figure 2.
Genotypic correlation coefficients for fruit traits of the 13 Platonia insignis genotypes collected from natural populations on the Brazilian Amazonian coast. Longitudinal fruit diameter (LFD), transversal fruit diameter (TFD), fruits pericarp thickness (FST), fruit color (FC), LFD/TFD ratio, fruit fresh mass (FFM), peel fresh mass (PFM), pulp fresh mass (PuFM), seed fresh mass (SFM), total number of parthenocarpy segments per fruit (NPSF), total number of seeds per fruit (NSF), peel yield (PY), pulp yield (PUY), seed yield (SY), soluble solids content (SSC), and titratable acidity (TTA). * p ≤ 0.05; ** p ≤ 0.01; and ns non-significant.
Figure 3.
Principal component analysis (PCA) of 16 morpho-agronomic traits in fruits of Platonia insignis from 13 genotypes sampled in natural Amazonian populations. (A) Biplot of the first two principal components (PC1 and PC2). Arrows indicate trait loadings, while points represent individual genotypes grouped into two clusters based on hierarchical clustering. Traits located farther from the origin exert a stronger influence on group separation. (B) Contribution of each trait to the variation explained by PC1. (C) Contribution of each trait to PC2, with peel fresh mass (PFM), peel yield (PY), and fruit color (FC) contributing most strongly. Trait acronyms: longitudinal fruit diameter (LFD), transversal fruit diameter (TFD), fruits pericarp thickness (FST), fruit color (FC), LFD/TFD ratio, fruit fresh mass (FFM), peel fresh mass (PFM), pulp fresh mass (PuFM), seed fresh mass (SFM), number of parthenocarpic segments (NPSF), number of seeds per fruit (NSF), peel yield (PY), pulp yield (PUY), seed yield (SY), soluble solids content (SSC), and titratable acidity (TTA).
Figure 3.
Principal component analysis (PCA) of 16 morpho-agronomic traits in fruits of Platonia insignis from 13 genotypes sampled in natural Amazonian populations. (A) Biplot of the first two principal components (PC1 and PC2). Arrows indicate trait loadings, while points represent individual genotypes grouped into two clusters based on hierarchical clustering. Traits located farther from the origin exert a stronger influence on group separation. (B) Contribution of each trait to the variation explained by PC1. (C) Contribution of each trait to PC2, with peel fresh mass (PFM), peel yield (PY), and fruit color (FC) contributing most strongly. Trait acronyms: longitudinal fruit diameter (LFD), transversal fruit diameter (TFD), fruits pericarp thickness (FST), fruit color (FC), LFD/TFD ratio, fruit fresh mass (FFM), peel fresh mass (PFM), pulp fresh mass (PuFM), seed fresh mass (SFM), number of parthenocarpic segments (NPSF), number of seeds per fruit (NSF), peel yield (PY), pulp yield (PUY), seed yield (SY), soluble solids content (SSC), and titratable acidity (TTA).
Figure 4.
Location of area where trees from natural populations of P. insignis were sampled, in Brazilian Amazon territory in the state of Maranhão (dark yellow). Municipalities: (1) São Luís, (2) Paço do Lumiar, (3) Santa Rita, (4) Alcântara, (5) Bequimão, (6) Pinheiro, (7) Nova Olinda, and (8) Codó.
Figure 4.
Location of area where trees from natural populations of P. insignis were sampled, in Brazilian Amazon territory in the state of Maranhão (dark yellow). Municipalities: (1) São Luís, (2) Paço do Lumiar, (3) Santa Rita, (4) Alcântara, (5) Bequimão, (6) Pinheiro, (7) Nova Olinda, and (8) Codó.
Table 1.
Estimates of genetic parameters for morpho-agronomic traits in Platonia insignis fruits from natural populations on the Brazilian Amazonian coast.
Table 1.
Estimates of genetic parameters for morpho-agronomic traits in Platonia insignis fruits from natural populations on the Brazilian Amazonian coast.
| Trait * | Parameter ** | |||||||
|---|---|---|---|---|---|---|---|---|
| Vg | Ve | Vf | h2g | h2mc | Acc | CVr | M | |
| LFD (mm) | 291.93 | 69.76 | 362.00 | 0.81 | 0.99 | 0.99 | 27.47 | 96.11 |
| TFD (mm) | 0.09 | 0.25 | 0.35 | 0.27 | 0.99 | 0.99 | 11.83 | 1.80 |
| FST (mm) | 55.61 | 41.35 | 97.02 | 0.57 | 0.99 | 0.99 | 24.02 | 82.90 |
| FC | 17.41 | 56.22 | 73.74 | 0.23 | 0.93 | 0.96 | 1.65 | 15.31 |
| LFD/TFD | 3.62 | 3.01 | 6.68 | 0.54 | 0.99 | 0.99 | 8.99 | 4.03 |
| FFM (g) | 5933.70 | 3090.13 | 9027.80 | 0.66 | 0.99 | 0.99 | 27.70 | 240.24 |
| PFM (g) | 154.24 | 488.83 | 644.42 | 0.24 | 0.99 | 0.99 | 8.90 | 64.56 |
| PuFM (g) | 0.52 | 0.77 | 1.30 | 0.40 | 0.99 | 0.99 | 18.63 | 2.91 |
| SFM (g) | 559.47 | 181.42 | 741.94 | 0.75 | 0.99 | 0.99 | 21.02 | 46.05 |
| NPSF (unit) | 6.67 | 17.72 | 24.42 | 0.27 | 0.99 | 0.99 | 11.21 | 11.57 |
| NSF (unit) | 0.33 | 0.56 | 0.90 | 0.37 | 0.99 | 0.99 | 17.24 | 2.19 |
| PY (%) | 27.72 | 84.51 | 112.75 | 0.24 | 0.93 | 0.97 | 1.67 | 22.60 |
| PUY (%) | 14.63 | 297.67 | 312.51 | 0.05 | 0.68 | 0.83 | 0.66 | 63.15 |
| SY (%) | 0.57 | 89.37 | 126.26 | 0.00 | 0.06 | 0.24 | 0.11 | 19.89 |
| SSC (°Brix) | 1.00 | 3.33 | 10.17 | 0.10 | 0.44 | 0.67 | 0.40 | 3.16 |
| TTA | 1.07 | 107.93 | 310.73 | 0.00 | 0.02 | 0.16 | 0.07 | 19.40 |
* Longitudinal fruit diameter (LFD), transversal fruit diameter (TFD), fruits pericarp thickness (FST), fruit color (FC), LFD/TFD ratio, fruit fresh mass (FFM), peel fresh mass (PFM), pulp fresh mass (PuFM), seed fresh mass (SFM), total number of parthenocarpy segments per fruit (NPSF), total number of seeds per fruit (NSF), peel yield (PY), pulp yield (PUY), seed yield (SY), soluble solids content (SSC), and titratable acidity (TTA). ** Genotypic variance between genotypes (Vg), residual variance (Ve), individual phenotypic variance (Vf), coefficient of relative variation (CVr), broad-sense heritability of individual plants (h2g), heritability on a mean basis of clones (h2mc), accuracy (Acc), and an overall average of the experiment (M).
Table 2.
Results of deviance analysis and likelihood ratio test (LRT) for the significance of genotypic variance (Vg) in morpho-agronomic traits of Platonia insignis fruits from natural populations along the Brazilian Amazonian coast.
Table 2.
Results of deviance analysis and likelihood ratio test (LRT) for the significance of genotypic variance (Vg) in morpho-agronomic traits of Platonia insignis fruits from natural populations along the Brazilian Amazonian coast.
| Trait | Deviance | LRT (Likelihood Ratio Test) |
|---|---|---|
| LFD (mm) | 1756.18 | 453.40 ** |
| TFD (mm) | −68.79 | 68.67 ** |
| FST (mm) | 1574.69 | 218.39 ** |
| FC | 1234.53 | 35.41 ** |
| LFD/TFD | 737.72 | 199.50 ** |
| FFM (g) | 2959.3 | 282.66 ** |
| PFM (g) | 2349.17 | 56.60 ** |
| PuFM (g) | 294.06 | 122.67 ** |
| SFM (g) | 2058.67 | 381.62 ** |
| NPSF (unit) | 1289.29 | 68.52 ** |
| NSF (unit) | 174.32 | 90.49 ** |
| PY (%) | 1359.89 | 42.11 ** |
| PUY (%) | 1654.63 | 2.98 ns |
| SY (%) | 1366.26 | 0.07 ns |
| SSC (°Brix) | 594.09 | 37.50 ** |
| TTA | 927.21 | 0.08 ns |
Longitudinal fruit diameter (LFD), transversal fruit diameter (TFD), fruits pericarp thickness (FST), fruit color (FC), LFD/TFD ratio, fruit fresh mass (FFM), peel fresh mass (PFM), pulp fresh mass (PuFM), seed fresh mass (SFM), total number of parthenocarpy segments per fruit (NPSF), total number of seeds per fruit (NSF), peel yield (PY), pulp yield (PUY), seed yield (SY), soluble solids content (SSC), and titratable acidity (TTA). Significance levels: p < 0.01 (**), ns = not significant.
Table 3.
Classification based on the sum of ranks for the top six Platonia insignis genotypes from natural populations on the Brazilian Amazonian coast.
Table 3.
Classification based on the sum of ranks for the top six Platonia insignis genotypes from natural populations on the Brazilian Amazonian coast.
| Trait | Genotype | ||||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| G1 | G2 | G3 | G4 | G5 | G6 | G7 | G8 | G9 | G10 | G11 | G12 | G13 | |
| FFM (+) | 13 | 12 | 9 | 4 | 2 | 8 | 5 | 10 | 1 | 6 | 11 | 3 | 7 |
| PFM (−) | 4 | 6 | 2 | 8 | 5 | 9 | 13 | 7 | 12 | 10 | 1 | 3 | 11 |
| SFM (−) | 2 | 1 | 3 | 9 | 12 | 6 | 7 | 8 | 10 | 11 | 4 | 13 | 5 |
| PuFM (+) | 2 | 3 | 8 | 7 | 12 | 4 | 1 | 6 | 10 | 9 | 5 | 13 | 11 |
| NSF (−) | 3 | 2 | 9 | 6 | 11 | 5 | 1 | 7 | 12 | 10 | 4 | 13 | 8 |
| NPSF (+) | 13 | 12 | 10 | 3 | 4 | 5 | 1 | 9 | 2 | 8 | 11 | 7 | 6 |
| FST (−) | 1 | 2 | 4 | 11 | 9 | 8 | 1 | 5 | 13 | 7 | 3 | 10 | 6 |
| PUY (+) | 2 | 10 | 11 | 12 | 2 | 5 | 4 | 8 | 1 | 7 | 13 | 9 | 3 |
| PY (−) | 12 | 8 | 3 | 5 | 4 | 7 | 6 | 2 | 1 | 10 | 9 | 11 | 13 |
| SY (−) | 8 | 4 | 1 | 2 | 6 | 3 | 9 | 5 | 10 | 11 | 12 | 13 | 7 |
| SST (+) | 12 | 13 | 11 | 10 | 8 | 7 | 5 | 6 | 9 | 2 | 4 | 1 | 3 |
| ATT (−) | 2 | 3 | 4 | 1 | 5 | 7 | 6 | 8 | 10 | 11 | 9 | 12 | 13 |
| Average rank | 6.17 | 6.33 | 6.25 | 6.50 | 6.67 | 6.17 | 4.92 | 6.75 | 7.58 | 8.50 | 7.17 | 9.00 | 7.75 |
Fruit fresh mass (FFM+), peel fresh mass (PFM−), seed fresh mass (SFM−), pulp fresh mass (PuFM+), number of seeds per fruit (NSF−), number of parthenocarpic segments per fruit (NPSF+), fruits pericarp thickness (FST−), pulp yield (PUY+), peel yield (PY−), seed yield (SY−), soluble solids content (SSC+), and titratable acidity (TTA−). Bolded average ranks highlight genotypes with the lowest overall values, indicating higher performance potential.
Table 4.
Geographic coordinates of the 13 Platonia insignis genotypes from natural populations on the Brazilian Amazonian coast.
Table 4.
Geographic coordinates of the 13 Platonia insignis genotypes from natural populations on the Brazilian Amazonian coast.
| Genotype | Municipality | Altitude | Latitude | Longitude |
|---|---|---|---|---|
| G1 | Alcântara | 515,510,403 | 2°32′2.34″ S | 44°38′23.61″ W |
| G2 | Bequimão | 233,110,114 | 2°30′2.06″ S | 44°44′39.56″ W |
| G3 | Bequimão | 223,636,639 | 2°30′2.65″ S | 44°52′48.15″ W |
| G4 | Bequimão | 221,215,877 | 2°30′1.78″ S | 44°52′47.23″ W |
| G5 | Codó | 634,813,685 | 4°27′39.93″ S | 43°47′7.22″ W |
| G6 | Codó | 73,011,913 | 4°27′42.23″ S | 43°47′15.48″ W |
| G7 | Paço do Lumiar | 246,098,218 | 2°31′55.82″ S | 44°10′32.97″ W |
| G8 | Nova Olinda | 581,240,947 | 2°41′21.30″ S | 45°42′32.90″ W |
| G9 | Pinheiros | 311,854,263 | 2°39′24.98″ S | 45°12′55.11″ W |
| G10 | Pinheiros | 204,556,364 | 2°25′21.78″ S | 45°8′46.12″ W |
| G11 | Pinheiros | 9,774,121 | 2°31′56.42″ S | 45°7′20.73″ W |
| G12 | São Luís | 489,158,026 | 2°36′2.48″ S | 44°12′ 44.43″ W |
| G13 | Santa Rita | 181,006,112 | 3°3′32.4″ S | 44°15′ 22.75″ W |
Table 5.
Classification of magnitudes of individual broad-sense heritability and selection accuracy used for estimating genetic parameters of Platonia insignis.
Table 5.
Classification of magnitudes of individual broad-sense heritability and selection accuracy used for estimating genetic parameters of Platonia insignis.
| Heritability Range (h2) | Heritability Classification | Accuracy Range (râa) | Accuracy Classification |
|---|---|---|---|
| 0.00–0.15 | Low | 0.10–0.40 | Low |
| 0.15–0.50 | Moderate | 0.40–0.70 | Moderate |
| ≥0.50 | High | ≥0.70 | High |
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Ribeiro, S.S.M.; Felipe, S.H.S.; Alves, G.L.; Rivas, P.M.S.; Henschel, J.M.; de Moraes, L.R.R.; Reis, L.C.F.; Araújo, J.R.G.; Pinheiro, M.V.M.; Batista, D.S.; et al. Genetic Variability, Broad-Sense Heritability, and Selection of Superior Genotypes for Fruit Improvement in Platonia insignis. Int. J. Plant Biol. 2025, 16, 108. https://doi.org/10.3390/ijpb16030108
AMA Style
Ribeiro SSM, Felipe SHS, Alves GL, Rivas PMS, Henschel JM, de Moraes LRR, Reis LCF, Araújo JRG, Pinheiro MVM, Batista DS, et al. Genetic Variability, Broad-Sense Heritability, and Selection of Superior Genotypes for Fruit Improvement in Platonia insignis. International Journal of Plant Biology. 2025; 16(3):108. https://doi.org/10.3390/ijpb16030108
Chicago/Turabian StyleRibeiro, Suzane Sá Matos, Sérgio Heitor Sousa Felipe, Givago Lopes Alves, Priscila Marlys Sá Rivas, Juliane Maciel Henschel, Lúcio Rafael Rocha de Moraes, Luís Carlos Ferreira Reis, José Ribamar Gusmão Araújo, Marcos Vinícius Marques Pinheiro, Diego Silva Batista, and et al. 2025. "Genetic Variability, Broad-Sense Heritability, and Selection of Superior Genotypes for Fruit Improvement in Platonia insignis" International Journal of Plant Biology 16, no. 3: 108. https://doi.org/10.3390/ijpb16030108
APA StyleRibeiro, S. S. M., Felipe, S. H. S., Alves, G. L., Rivas, P. M. S., Henschel, J. M., de Moraes, L. R. R., Reis, L. C. F., Araújo, J. R. G., Pinheiro, M. V. M., Batista, D. S., & Corrêa, T. R. (2025). Genetic Variability, Broad-Sense Heritability, and Selection of Superior Genotypes for Fruit Improvement in Platonia insignis. International Journal of Plant Biology, 16(3), 108. https://doi.org/10.3390/ijpb16030108
