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Research Article

Phenotypic Characteristics of Quantitative Traits in an Everbearing Strawberry F₁ Population

Iyeong Choi1, Young Hun Lee2,*
Plant Breeding and Biotechnology 2025;13:229-242.
Published online: October 28, 2025

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*Corresponding to Young Hun Lee TEL. +82-33-640-2913 E-mail. hunly@gwnu.ac.kr
• Received: September 26, 2025   • Revised: September 29, 2025   • Accepted: September 30, 2025

Copyright © 2025 by the Korean Society of Breeding Science

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  • Everbearing strawberry cultivars provide fruit during the summer–fall period when June-bearing strawberries are unavailable, but their breeding progress has been constrained by complex trait interactions. To characterize segregation patterns and evaluate phenotypic diversity, we developed an F₁ population from a cross between two everbearing cultivars, ‘Charlotte’ and ‘Flamenco’. Twenty selected progenies were evaluated for 30 quantitative traits encompassing vegetative vigor, inflorescence structure, fruit morphology, firmness, and biochemical composition. Substantial variation was observed among lines, with several individuals exceeding parental performance for key traits such as fruit size, soluble solids content, and sucrose accumulation. Principal component analysis revealed three major axes of variation: fruit composition (sugars and acids), vegetative vigor, and fruit size and morphology. K-means clustering grouped the progeny into three phenotypic classes, representing high-sugar, large-fruited, or vigorous growth types. Notably, some lines combined favorable attributes across classes, such as high sweetness and large fruit, indicating the potential to overcome typical trade-offs between yield and quality. These findings provide a practical framework for breeding selection and highlight superior progeny as immediate candidates for clonal advancement or as parents in future crossing. The results also establish a foundation for integrating phenotypic classifications with molecular tools to accelerate the development of high-value everbearing strawberries.
  • Key words: Strawberry, Everbearing, F₁ population, Phenotypic variation, Quantitative traits
Cultivated strawberry (Fragaria × ananassa Duch.) is a globally important fruit crop with annual production exceeding 9 million tonnes worldwide (Kouloumprouka Zacharaki et al. 2024). In Korea, most commercial production relies on June-bearing cultivars grown in greenhouses from winter to spring. Since the early 2000s, however, growers in high-altitude regions above 800 m have adopted everbearing cultivars that initiate flowers under long-day, high-temperature conditions. This enables fruit harvest during the summer–fall period when June-bearing strawberries are unavailable and provides a premium fresh product in domestic and export markets.
Flowering physiology differs fundamentally between June-bearing and everbearing cultivars. June-bearing types initiate flower buds under short days, whereas everbearing types maintain flowering under long days and warm temperatures. The major locus PFRU interacts with FaTFL1, a floral repressor integrating photoperiod and temperature signals (Hytönen et al. 2020; Rantanen et al. 2015). While flowering regulation has been studied extensively in June-bearing cultivars, research focusing on everbearing cultivars remains relatively limited, making it more difficult to improve yield stability and fruit quality in breeding programs.
Because of the allo-octoploid genome and high heterozygosity, F1 progenies from elite crosses segregate broadly for quantitative traits, and even first-generation seedlings may display transgressive segregation for fruit weight, firmness, soluble solids content, acidity, or yield components (Lerceteau-Köhler et al. 2012; Perrotte et al. 2016; Samad et al. 2017; Zorrilla-Fontanesi et al. 2011). However, most previous genetic studies have examined traits in isolation or have used mixed photoperiod populations such as June-bearing × everbearing crosses. Integrated multivariate analyses of trait segregation within everbearing × everbearing populations remain scarce.
Therefore, the objectives of this study were (1) to characterize segregation patterns across vegetative, floral, and fruit traits in an F1 population derived from the everbearing cultivars ‘Charlotte’ and ‘Flamenco’, and (2) to analyze inter-trait correlations using multivariate methods to inform ideotype-based breeding strategies.
Plant materials and F₁ population development
The F1 population was generated by crossing the everbearing cultivar ‘Charlotte’ (France, released 1995) with ‘Flamenco’ (UK, East Malling Research, released 2002). ‘Charlotte’ was chosen as the female parent for its superior fruit shape and quality and is the most widely grown everbearing cultivar in the highland region of Daegwallyeong, Korea (37.6811°N, 128.7191°E; 781.0 m elevation). ‘Flamenco’, used as the male parent, is noted for its high fruit firmness and low incidence of malformed fruit under high-temperature conditions. Hand pollinations were conducted from September to November 2013, yielding approximately 300 F1 seeds from the cross.
Harvested seeds were cold-stratified at 2℃ for 8 weeks to break dormancy, then surface-sterilized in 2% sodium hypochlorite (NaOCl) for 8 hours. Sterilized seeds were sown on paper towels moistened with distilled water in sealed plastic bags and allowed to germinate for 30 days, producing 221 seedlings. Seedlings were transplanted into 32-cell trays filled with a commercial soilless substrate (Nongwoo Bio Co., Ltd., Suwon, Republic of Korea) and grown in a glasshouse at Gangneung-Wonju National University from December 2013 to June 2014. Thereafter, 158 healthy plantlets were transplanted to a greenhouse at the university’s research farm (37.7714°N, 128.8664°E; 38.5 m elevation) and arranged in a completely randomized layout under standard cultivation conditions. From this population, 20 F1 lines exhibiting superior horticultural performance were selected for detailed quantitative evaluation (Table 1). The pedigree of the ‘Charlotte’ × ‘Flamenco’ cross is illustrated in Fig. 1.
Trait measurements
Thirty horticultural traits were assessed in the selected F1 lines (Table 1), comprising 19 vegetative/inflorescence traits and 11 fruit-related traits. Unless otherwise noted, each trait was measured with at least three replicates per line (e.g., three fruits or three measurements per genotype), and the line mean was used for analysis (Table 2). Qualitative horticultural descriptors (e.g., growth habit, leaf density, fruit shape category) were recorded according to the International Union for the Protection of New Varieties of Plants (UPOV) strawberry guidelines.
Vegetative and inflorescence traits included plant height, plant spread (canopy diameter), terminal leaflet length and width, petiole length, leaf area, leaf color (SPAD), the node position of the first inflorescence, number of flowers per inflorescence, peduncle length, and peduncle angle. Leaf dimensions and plant height were recorded over a 60-day active growth period using a ruler and vernier calipers; leaf area on fully expanded trifoliates was measured with a leaf area meter (GA-5, Tokyo Photoelectric Co., Tokyo, Japan) by summing the area of the three leaflets. Leaf color (chlorophyll content) was quantified as SPAD values using a handheld chlorophyll meter (SPAD-502, Konica Minolta, Tokyo, Japan). Growth habit, leaf density, and overall vigor were visually scored following UPOV descriptors. The node of first inflorescence was defined as the number of nodes on the crown before the first flower truss; flower number was recorded at first bloom. Peduncle (inflorescence stalk) orientation was noted qualitatively (erect, semi-erect, or drooping relative to horizontal).
Fruit morphology and firmness were evaluated on representative ripe fruits. Fruit length (proximal–distal) and width (equatorial diameter) were measured with vernier calipers. Fruit firmness (penetration force) was determined at the equatorial region using a texture analyzer (EZ Test-100N, Shimadzu Co., Kyoto, Japan) fitted with a 5 mm-diameter cylindrical probe moving at a constant speed of 120 mm per minute; higher force readings indicate firmer flesh.
Primary fruit composition was analyzed by high-performance liquid chromatography (HPLC). Approximately 2 g of fruit mesocarp (flesh) per line was extracted in 40 mL of 90% methanol containing 5 mM butylated hydroxytoluene (BHT) as an antioxidant. The homogenate was centrifuged, and the supernatant was filtered through a 0.45 µm syringe filter before HPLC injection (LC-10AT system, Shimadzu Co., Kyoto, Japan). Soluble sugars (glucose, fructose, and sucrose) were separated on a Rezex RCM-Monosaccharide column (Phenomenex, Torrance, CA, USA) with deionized water as the mobile phase and detected with a refractive index detector (RID-10A, Shimadzu). Organic acids (citric acid and malic acid) were separated on an Alltech IOA-1000 organic acid column (Grace-Alltech, Deerfield, IL, USA) with 5 mM H₂SO₄ as the mobile phase and quantified by UV absorbance detection.
Statistical analysis
To summarize variation in the F1 population, principal component analysis (PCA) was performed on the correlation matrix of the 30 quantitative traits; scatter plots of line scores on the first principal components were used to visualize patterns of variation and clustering among genotypes. K-means clustering was then applied to group the 20 F1 lines into clusters with similar multi-trait profiles, with the number of clusters chosen by examining within-cluster variance and biological interpretability. For trait-wise comparisons, one-way analysis of variance (ANOVA) was conducted for each quantitative trait. When ANOVA indicated significant differences, pairwise comparisons of line means were carried out using Scheffé’s multiple range test at the 5% significance level (p < 0.05). All statistical analyses were implemented using IBM SPSS Statistics version 21.0 (IBM Corp., Armonk, NY, USA).
Vegetative and floral characteristics
Plant height, plant spread, growth habit, leaf density, and plant vigor were evaluated for 20 lines (Table 1). GWEBS-69 had the greatest plant height, whereas GWEBS-18 and GWEBS-152 were the shortest. Plant spread was also greatest in GWEBS-69 and smallest in GWEBS-18. Growth habit was mostly semi-erect, similar to the parents, although GWEBS-82 was classified as erect. Leaf density and plant vigor were intermediate across all lines. These results suggest that plant height and spread are strongly influenced by environmental variation, whereas growth habit, leaf density, and vigor are predominantly governed by genetic factors.
Terminal leaflet length, width, length-to-width ratio, and shape were also recorded (Table 2). GWEBS-69 had the greatest leaflet length, while GWEBS-147 had the smallest; leaflet width was also greatest in GWEBS-69 and smallest in GWEBS-99. The length-to-width ratio was highest in GWEBS-99 and lowest in GWEBS-66; however, this ratio did not correlate with leaflet shape. Representative leaflet and floral variation across lines are shown in Fig. 2. For leaf color, leaf area, and petiole length, GWEBS-05 showed the highest chlorophyll content, while GWEBS-43 was the lowest. Leaf area was greatest in GWEBS-12 and smallest in GWEBS-99. Petiole length was greatest in ‘Flamenco’ and, among the progeny, in GWEBS-43; GWEBS-12 had the shortest petiole. Overall, ‘Charlotte’ and ‘Flamenco’ exhibited longer petioles than the hybrid lines (Table 2).
Node position of the first inflorescence, number of flowers, peduncle length, and peduncle angle were evaluated (Table 3). The first inflorescence appeared at the fewest nodes in GWEBS-66 and at the most nodes in GWEBS-18. The greatest flower number occurred in GWEBS-12, and the fewest in GWEBS-127. ‘Charlotte’ had the longest peduncle; among lines, GWEBS-18 was the longest and GWEBS-147 the shortest. Most lines showed an erect or semi-erect peduncle angle, although downward-facing inflorescences were observed in some lines. Representative flower morphologies of selected lines are presented in Fig. 3.
Fruit characteristics
Fruit morphological traits varied widely among ‘Charlotte’ × ‘Flamenco’ progeny (Table 4). Mean fruit weight ranged from the largest in GWEBS-12 to the smallest in GWEBS-127. Most lines produced flat or round berries similar to the parental shapes, although a few exhibited more elongated, oblong types. For example, GWEBS-66 produced oblong fruit not prominent in either parent. The calyx-end diameter also showed segregation: it was greatest in GWEBS-66 and smallest in GWEBS-43. Internal fruit morphology differed among lines as well: hollow cores were generally moderate, but were prominent in GWEBS-05 and GWEBS-90. Representative external and cross-sectional images of fruit types are shown in Fig. 4.
Substantial variation was also observed in fruit biochemical composition (Table 4). Soluble solids content (°Brix) ranged widely, reflecting differences in sweetness. Individual sugars (sucrose, glucose, fructose) varied markedly, as did citric and malic acid contents. Notably, some lines combined high sucrose concentration with high °Brix, producing exceptionally sweet fruit. In contrast, others had reduced citric acid proportion, yielding a milder taste. Certain progeny matched or exceeded the sucrose levels of ‘Charlotte’ while also accumulating high total sugars, and some had significantly lower citric acid than either parent, giving a smoother flavor profile. These results demonstrate that desirable flavor attributes—high sweetness with balanced acidity—can be identified within this single-cross population.
Statistical analysis
Principal component analysis (PCA) extracted three components with eigenvalues ≥ 1, together explaining ~47% of the total variance. PC1 was strongly associated with fruit compositional traits, including fructose, glucose, sucrose, soluble solids content, and sepal width. PC2 correlated with vegetative traits such as terminal leaflet dimensions, peduncle length, plant height, and canopy spread. PC3 was primarily related to fruit size and form, including fruit width, fruit length, and sepal length/width ratio. Comparable PCA-based differentiation of strawberry germplasm has been reported previously in Korea, supporting the robustness of multivariate approaches for trait classification {Kim et al. 2009 #14}.
Scatter plots of PCA scores revealed clear separation among lines (Fig. 5). Lines on the positive end of PC1 (e.g., GWEBS-66 and GWEBS-130) formed a cluster characterized by high sugars and °Brix, whereas lines on the negative end were low in sugar. Along PC2, lines with greater vegetative biomass occupied the upper region, while less vigorous types were positioned lower. PC3 discriminated fruit morphology, with certain lines exhibiting large or elongated fruit clearly separating from others.
K-means clustering grouped the 20 F1 lines into three distinct clusters (Fig. 6), hereafter referred to as Cluster 1, Cluster 2, and Cluster 3. Cluster 1 comprised lines with superior vegetative growth (high PC2 scores), Cluster 2 included lines characterized by pronounced fruit size and morphology (high PC3 scores), and Cluster 3 consisted of lines with superior fruit composition, particularly high sugar content and soluble solids (high PC1 scores). This clustering aligned closely with the PCA axes and indicates that major phenotypic axes correspond to distinct horticultural ideotypes.
To validate the clusterwise separations observed in PCA, ANOVA was conducted on principal component scores. Significant differences were detected among the three clusters for all principal components (p < 0.05). Cluster 3 showed the highest mean PC1, consistent with its high-sugar profile. Cluster 1 had the highest mean PC2, corresponding to vigorous vegetative growth. Cluster 2 showed the highest mean PC3, reflecting larger fruit size and distinctive morphology. Together, PCA, clustering, and ANOVA provide a coherent statistical framework for distinguishing ideotype groups within the population.
Phenotypic variation and trait associations
The F1 population derived from ‘Charlotte’ × ‘Flamenco’ displayed broad segregation across vegetative, floral, and fruit traits (Tables 1–4; Figs. 2–4). This variability is consistent with the high heterozygosity of octoploid strawberry and the frequent occurrence of transgressive phenotypes reported in similar crosses (Lerceteau-Köhler et al. 2012; Zorrilla-Fontanesi et al. 2011). Such variation is also comparable to field observations from highland summer culture in Korea, where cultivar characteristics and crown management influenced yield stability of everbearing strawberries{Hwang et al. 2013 #17}.
Vegetative traits such as plant height, canopy spread, and petiole length varied substantially (Table 1), with GWEBS-69 exhibiting the tallest plants and widest spread, whereas GWEBS-18 and GWEBS-152 were compact. These results highlight that basic vigor traits segregate widely, providing potential for selecting genotypes adapted to different production systems.
Floral characteristics also showed clear segregation (Table 3; Fig. 3). For example, node position of the first inflorescence differed between lines, ranging from GWEBS-66 with low node numbers to GWEBS-18 with high node numbers. Peduncle orientation varied from erect to drooping, which can directly affect pollination efficiency and fruit exposure. Such traits are underpinned by flowering regulatory networks involving TFL1/FT/SOC1 modules (Hytönen et al. 2020; Rantanen et al. 2015). These pathways integrate photoperiod and temperature signals, thereby linking observed floral variation to physiological mechanisms of flowering control. The correlation between vegetative vigor (PC2) and fruit set/size (PC3) (Fig. 5) likely reflects this shared regulatory background. Previous studies have shown that flowering time and axillary bud fate are highly environment-sensitive, with QTL identified for these traits in both diploid and octoploid strawberry (Andrés et al. 2021; Prohaska et al. 2024; Samad et al. 2017). Our findings reinforce that floral variation is not only a morphological outcome but also an axis of adaptive and reproductive strategies in everbearing strawberry.
Fruit morphology and composition showed the widest range of segregation (Table 4; Fig. 4). Large-fruited types such as GWEBS-12 contrasted with small-fruited progeny like GWEBS-127. Novel oblong fruit shapes appeared in lines such as GWEBS-66, not evident in the parents, while internal structures such as prominent hollow cores were observed in GWEBS-05 and GWEBS-90. Biochemical composition further diversified the population: some lines combined high °Brix and sucrose (e.g., GWEBS-66, GWEBS-130), while others exhibited lower citric acid levels, conferring milder flavor. These findings demonstrate that transgressive combinations can produce fruits surpassing parental quality standards, breaking the commonly observed trade-off between sweetness and yield (Cockerton et al. 2021).
Multivariate analysis summarized these complex traits into three orthogonal components: PC1 (fruit composition), PC2 (vegetative vigor), and PC3 (fruit size and morphology) (Fig. 5). K-means clustering grouped the 20 lines into three major classes (Fig. 6): high-sugar, large-fruited, and vigorous types. This classification provides a practical phenotypic grouping that aligns with horticultural targets in everbearing strawberry improvement.
Breeding implications and future perspectives
The phenotypic groups identified here provide immediate implications for strawberry breeding. High-sugar lines such as GWEBS-66 and GWEBS-130 (Table 4) represent elite flavor-oriented materials. Large-fruited progeny such as GWEBS-12 (Fig. 4) are valuable for market-preferred fruit size, while vigorous types such as GWEBS-69 (Table 1) offer productivity advantages in high-density systems. These superior lines can be advanced clonally as potential cultivars and also serve as diverse parents for targeted crossing.
Several progeny demonstrated transgressive performance that exceeded both parents, such as GWEBS-12 combining very large fruit with high °Brix, and GWEBS-05 and GWEBS-90 exhibiting unique internal morphologies. These lines exemplify how single-cross populations can generate novel phenotypes that expand the breeding pool.
Planned inter-group crosses are particularly valuable: for instance, crossing high-sugar lines (Cluster 3, Fig. 6) with large-fruit lines (Cluster 2, Fig. 6) could yield progeny that integrate both sweetness and marketable size. This approach takes advantage of the independent segregation patterns revealed by PCA and clustering, creating opportunities to recombine favorable alleles and overcome trade-offs traditionally limiting improvement.
Molecular tools provide complementary support. Stable QTL for fruit shape and firmness in octoploid strawberry {Rey-Serra et al. 2021 #11} correspond to traits captured in PC3, making marker-assisted selection feasible. Genomic prediction, when coupled with high-throughput phenotyping of fruit composition traits such as sugars and acids (PC1), can accelerate selection for polygenic traits like sweetness and yield {Cockerton et al. 2021 #10}. Moreover, the development of octoploid SNP marker systems provides a robust genomic resource that, when integrated with phenotypic grouping, further enhances the precision and efficiency of breeding strategies {Lee et al. 2017 #12}.
A limitation of this study is that the evaluation was conducted in a single environment. Since flowering, vigor, and fruit quality are all subject to genotype × environment interactions, validation across multiple seasons and sites is essential. Multi-environment trials will confirm the stability of the phenotypic groups identified here and the robustness of their breeding value. Ultimately, combining phenotypic group-based strategies with molecular markers, genomic prediction, and precision cultivation will accelerate genetic gain and facilitate the release of high-value everbearing cultivars that meet both grower productivity needs and consumer preferences.
This research was supported by the Regional Innovation System & Education(RISE) Glocal University 30 Project program through the Gangwon RISE Center, funded by the Ministry of Education(MOE) and the Gangwon State(G.S.), Republic of Korea.(2025-RISE-10-004)
Fig. 1
Pedigree diagram of everbearing strawberry cultivars used in the cross ‘Charlotte’ × ‘Flamenco’.
pbb-13-229-f1.jpg
Fig. 2
Morphological variation in flowers and fruits of the F1 progeny from ‘Charlotte’ × ‘Flamenco’. Representative images illustrate floral shapes and fruit types compared with parental cultivars.
pbb-13-229-f2.jpg
Fig. 3
Principal component analysis (PCA) of 30 quantitative traits in the F1 progeny from ‘Charlotte’ × ‘Flamenco’. Score plots show distribution along the first three components.
pbb-13-229-f3.jpg
Fig. 4
Clustering of 20 F1 lines from ‘Charlotte’ × ‘Flamenco’ based on PCA scores. Three phenotypic groups were identified, corresponding to fruit composition, vegetative vigor, and fruit size.
pbb-13-229-f4.jpg
Fig. 5
Variation in fruit internal morphology among selected F1 lines compared with parental cultivars. Cross-sections highlight presence of hollow core and pith development.
pbb-13-229-f5.jpg
Fig. 6
Representative fruits of high-sugar, large-fruit, and vigorous-growth groups identified in the F1 population.
pbb-13-229-f6.jpg
Table 1
Plant characteristics of ever-bearing strawberry crossed lines.
Table 1
Line Plant height(cm) Plant spread(cm) Plant habit Plant density
of foliage		 Plant vigor
GWEBS-05 19.5 29.5 semi-upright M M
GWEBS-08 16.7 29.3 semi-upright M M
GWEBS-12 16.5 33.5 semi-upright M M
GWEBS-18 14.3 21.7 semi-upright M M
GWEBS-29 17.8 25.8 semi-upright M M
GWEBS-43 18.5 30.6 semi-upright M M
GWEBS-61 15.3 24.3 semi-upright M M
GWEBS-66 16.8 29.5 semi-upright M M
GWEBS-69 21.3 35.9 semi-upright M M
GWEBS-82 19.2 23.1 upright M M
GWEBS-90 20.5 26.8 semi-upright M M
GWEBS-99 15.2 25.7 semi-upright M M
GWEBS-111 18.9 34.7 semi-upright M M
GWEBS-116 20.5 28.5 semi-upright M M
GWEBS-120 17.1 28.9 semi-upright M M
GWEBS-127 17.5 30.9 semi-upright M M
GWEBS-128 17.2 30.5 semi-upright M M
GWEBS-130 14.5 29.8 semi-upright M M
GWEBS-147 15.7 26.7 semi-upright M M
GWEBS-152 14.2 29.1 semi-upright M M
Charlotte 19.8 34.6 semi-upright M M
Flamenco 18.1 30.33 semi-upright M M
Table 2
Foliage characteristics of ever-bearing strawberry crossed lines.
Table 2
Line Terminal leaflat length(mm) Terminal leaflat width
(mm)		 Terminal W/L ratio Terminal leaflet shape of base Leaf color SPAD value Leaf area
(mm2)		 Petiole length
(mm)
GWEBS-05 65.7 64.5 1.02 acute 45.6 66.9 126.7
GWEBS-08 72.3 72.1 1 obtuse 39.9 109.4 147.6
GWEBS-12 75.9 75.6 1 obtuse 43.5 119.8 86.5
GWEBS-18 53.7 53 1.01 obtuse 38 70.5 99
GWEBS-29 66.9 73.9 0.91 obtuse 38.8 72.7 117.7
GWEBS-43 57.2 58.3 0.98 obtuse 29.9 84.9 156.2
GWEBS-61 70.2 69.9 1 obtuse 40.5 110.9 97.3
GWEBS-66 64 73.5 0.87 rounded 38.8 68.3 95.2
GWEBS-69 85.7 87.1 0.98 acute 37.2 68.9 125.6
GWEBS-82 61.8 58 1.07 rounded 38.1 97.7 122.4
GWEBS-90 67.6 55.1 1.23 acute 37.3 106.4 130.6
GWEBS-99 70.6 51.4 1.37 obtuse 33.9 58.7 89.4
GWEBS-111 74.1 77.5 0.96 obtuse 36.4 107.5 108.3
GWEBS-116 75.6 64.2 1.18 obtuse 37.1 72.3 139.2
GWEBS-120 64.8 59.4 1.09 obtuse 39.9 59.7 114.7
GWEBS-127 71.7 67.8 1.06 acute 38.7 63 132.6
GWEBS-128 67.8 60.4 1.12 obtuse 40.3 79 103.4
GWEBS-130 66.2 63.5 1.04 obtuse 39.8 86.2 106.6
GWEBS-147 52 58.6 0.89 obtuse 42.8 65.4 113.2
GWEBS-152 62.1 62 1 obtuse 45 109.3 93.4
Charlotte 73.7 74.4 0.99 obtuse 43.5 107.7 142.3
Flamenco 81.8 70.2 1.17 obtuse 40.4 74.7 158.2
Table 3
Flower characteristics of ever-bearing strawberry crossed lines.
Table 3
Lines Calyx diameter (mm) Sepal length (mm) Sepal width (mm) Sepal W/L ratio Relation between flower size and calyx Petal length (mm) Petal width (mm) Petal L/W ratio No. of petal No. of flower Flower diameter (mm) Peduncle length (cm) Flower arrangement of petal
GWEBS-05 9.2 13.2 8.4 1.56 large 8.9 8.6 1.04 6.2 8.3 19 17.1 Overlapping
GWEBS-08 9.4 9.9 5.9 1.67 same 9.1 10.6 0.86 6 9.7 24.4 14.8 Touching
GWEBS-12 12.7 12.7 7.1 1.79 large 10 12.4 0.81 6 9.7 21.3 16.6 Overlapping
GWEBS-18 12.2 9.5 5.4 1.76 large 6.8 8 0.86 5.4 10 22.5 13.4 Touching
GWEBS-29 9.8 10.2 5.6 1.82 large 8 9.3 0.86 5.4 8.7 19.6 18.7 Overlapping
GWEBS-43 11.4 8.8 5.3 1.65 large 6.3 7.6 0.83 6 10.3 13.8 16.9 Overlapping
GWEBS-61 9.7 17.4 9.1 1.9 large 8.5 9.5 0.89 5 9.7 24.8 14.6 Overlapping
GWEBS-66 12.2 17.6 11.7 1.5 large 10.6 11.3 0.93 6 10 24.5 19.2 Overlapping
GWEBS-69 11.9 14.6 8.8 1.65 large 8.2 7.4 1.11 6 9.7 26.5 20.5 Free
GWEBS-82 12.1 11.1 9.6 1.16 large 7.2 7.4 0.98 6 10 30.3 16.2 Overlapping
GWEBS-90 9.2 9 6.2 1.44 large 7.7 7.5 1.03 6 9.7 19.4 15.4 Overlapping
GWEBS-99 8.6 7.8 5.1 1.54 same 7.8 7.8 0.99 5 11.3 17.8 16.8 Overlapping
GWEBS-111 9.3 11 7 1.57 large 6.5 6.6 0.98 6 11.3 19.5 16.4 Overlapping
GWEBS-116 10.5 15.9 8.6 1.85 large 8.9 9.3 0.96 5.8 11.7 26.4 17.4 Overlapping
GWEBS-120 10.2 11.9 8.3 1.44 large 7.7 7.6 1.02 5.4 9.3 22.9 14.2 Overlapping
GWEBS-127 11 10.5 5.4 1.95 large 7.2 8 0.9 5 11.3 23.4 18.5 Overlapping
GWEBS-128 11 14.4 7.7 1.88 large 7.9 7.9 1 5.4 11.7 20.9 15.4 Overlapping
GWEBS-130 9.2 14.9 9.3 1.61 large 8.5 7.1 1.19 5 8.3 19.8 16.3 Free
GWEBS-147 8.6 7.3 4.7 1.55 large 6.6 6.3 1.04 5.4 9.7 22.6 15.8 Overlapping
GWEBS-152 9.7 8.9 5.1 1.76 same 8.6 9.5 0.91 6 11 22.4 15.7 Overlapping
Charlotte 11.7 11.7 5.7 2.05 large 9 9.4 0.96 5.2 11.7 27.5 20.2 Free
Flamenco 12 13.7 6.4 2.14 large 8.3 8 1.04 5.8 11 28.2 19.5 overlapping
Table 4
Fruit characteristics of ever-bearing strawberry crossed lines.
Table 4
Lines Fruit length (mm) Fruit width (mm) Fruit L/W ratio Fruit firmness (N/5 mm) Fruit weight (g) Fruit shape Fruit position of achenes Fruit position of calyx attachment Fruit attitude of sepals Sucrose Glucose Fructose citric acid Malic acid Soluble solids (°Brix)
GWEBS-05 23.7 22.4 0.95 4.21 8.6 reniform level with surface inserted outwards 8.54 5.03 19.96 7.86 0.94 7.4
GWEBS-08 31.1 29.3 0.94 5.07 7.6 ovoid level with surface level with fruit outwards 19.19 20.38 40.7 5.84 5.13 7.2
GWEBS-12 28.2 27.6 0.98 3.22 15.7 obloid level with surface inserted outwards 16.23 13.29 34.49 6.33 4.92 9.5
GWEBS-18 30.8 27.7 0.9 4.66 14.1 conical above surface inserted outwards 6.57 9.85 38.73 4.74 6.36 7.5
GWEBS-29 27.7 27.1 0.98 5.73 9.4 conical level with surface level with fruit upwards 14.4 15.02 35.87 6.75 1.52 8.2
GWEBS-43 19.7 18.7 0.95 5.03 10.8 conical above surface level with fruit upwards 7.76 7.53 25.14 6.11 0.12 6.8
GWEBS-61 29.8 26.8 0.9 3.41 8.9 ovoid above surface raised outwards 13.91 17.01 76.7 6.61 3.02 10.6
GWEBS-66 28.1 22.2 0.79 3.92 12.5 globose above surface level with fruit outwards 22.25 24.73 50.84 6.07 0.93 8.8
GWEBS-69 24.9 24.7 0.99 3.29 10.4 globose level with surface level with fruit upwards 1.08 1.25 21.24 6.45 4.03 6.5
GWEBS-82 25 21.7 0.87 3.02 11.8 conical level with surface level with fruit upwards 20.01 14.01 37.18 6.79 1.04 8.6
GWEBS-90 25.3 24.6 0.97 4.91 8.6 conical level with surface level with fruit upwards 15.72 11.09 31.14 9.2 2.73 8.8
GWEBS-99 20.2 19.7 0.98 6.01 7.9 conical below surface level with fruit upwards 5.16 3.08 11.65 5.35 3.52 6.7
GWEBS-111 19.8 18.7 0.94 5.24 7 globose level with surface level with fruit outwards 21.97 11.16 35.03 6.76 1.82 8.9
GWEBS-116 25.4 27.1 1.07 3.75 10.7 wedged level with surface level with fruit outwards 9.95 8.05 28.31 8.4 2.82 7.4
GWEBS-120 26.5 24.9 0.94 3.31 10.8 reniform below surface level with fruit outwards 2.11 2.74 21.6 5.48 1.62 8.2
GWEBS-127 32.2 29.4 0.91 5.59 12.4 conical above surface level with fruit outwards 6.55 2.16 15.33 6.27 4.37 8.9
GWEBS-128 30.7 28.3 0.92 3.75 11.4 conical level with surface level with fruit upwards 4.91 13.28 36.46 4.02 0.93 8.7
GWEBS-130 33 30.1 0.91 3.74 7.9 cordate level with surface level with fruit upwards 23.72 16.74 32.87 6.27 4.28 10
GWEBS-147 25 21.5 0.86 4.69 6.7 ovoid level with surface level with fruit outwards 27.29 19.97 46.18 7.24 3.05 7.6
GWEBS-152 25.5 25.1 0.98 4.64 9.3 conical level with surface level with fruit outwards 8.96 7.02 21.5 5.7 0.02 6.9
Charlotte 31 32.4 1.05 4.3 10.7 conical level with surface level with fruit upwards 8.48 19.74 44.67 5.02 0.83 8.2
Flamenco 34.7 31.8 0.92 4.02 11.2 conical level with surface level with fruit upwards 18.77 17.05 36.62 5.71 5.42 7.7
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Phenotypic Characteristics of Quantitative Traits in an Everbearing Strawberry F₁ Population
Plant Breed. Biotech.. 2025;13:229-242.   Published online October 28, 2025
DOI: https://doi.org/10.9787/PBB.2025.13.229
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Phenotypic Characteristics of Quantitative Traits in an Everbearing Strawberry F₁ Population
Plant Breed. Biotech.. 2025;13:229-242.   Published online October 28, 2025
DOI: https://doi.org/10.9787/PBB.2025.13.229
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Phenotypic Characteristics of Quantitative Traits in an Everbearing Strawberry F₁ Population
Image Image Image Image Image Image
Fig. 1 Pedigree diagram of everbearing strawberry cultivars used in the cross ‘Charlotte’ × ‘Flamenco’.
Fig. 2 Morphological variation in flowers and fruits of the F1 progeny from ‘Charlotte’ × ‘Flamenco’. Representative images illustrate floral shapes and fruit types compared with parental cultivars.
Fig. 3 Principal component analysis (PCA) of 30 quantitative traits in the F1 progeny from ‘Charlotte’ × ‘Flamenco’. Score plots show distribution along the first three components.
Fig. 4 Clustering of 20 F1 lines from ‘Charlotte’ × ‘Flamenco’ based on PCA scores. Three phenotypic groups were identified, corresponding to fruit composition, vegetative vigor, and fruit size.
Fig. 5 Variation in fruit internal morphology among selected F1 lines compared with parental cultivars. Cross-sections highlight presence of hollow core and pith development.
Fig. 6 Representative fruits of high-sugar, large-fruit, and vigorous-growth groups identified in the F1 population.

Fig. 1

Fig. 2

Fig. 3

Fig. 4

Fig. 5

Fig. 6

Phenotypic Characteristics of Quantitative Traits in an Everbearing Strawberry F₁ Population

Plant characteristics of ever-bearing strawberry crossed lines.

Line Plant height(cm) Plant spread(cm) Plant habit Plant density
of foliage		 Plant vigor
GWEBS-05 19.5 29.5 semi-upright M M
GWEBS-08 16.7 29.3 semi-upright M M
GWEBS-12 16.5 33.5 semi-upright M M
GWEBS-18 14.3 21.7 semi-upright M M
GWEBS-29 17.8 25.8 semi-upright M M
GWEBS-43 18.5 30.6 semi-upright M M
GWEBS-61 15.3 24.3 semi-upright M M
GWEBS-66 16.8 29.5 semi-upright M M
GWEBS-69 21.3 35.9 semi-upright M M
GWEBS-82 19.2 23.1 upright M M
GWEBS-90 20.5 26.8 semi-upright M M
GWEBS-99 15.2 25.7 semi-upright M M
GWEBS-111 18.9 34.7 semi-upright M M
GWEBS-116 20.5 28.5 semi-upright M M
GWEBS-120 17.1 28.9 semi-upright M M
GWEBS-127 17.5 30.9 semi-upright M M
GWEBS-128 17.2 30.5 semi-upright M M
GWEBS-130 14.5 29.8 semi-upright M M
GWEBS-147 15.7 26.7 semi-upright M M
GWEBS-152 14.2 29.1 semi-upright M M
Charlotte 19.8 34.6 semi-upright M M
Flamenco 18.1 30.33 semi-upright M M

Foliage characteristics of ever-bearing strawberry crossed lines.

Line Terminal leaflat length(mm) Terminal leaflat width
(mm)		 Terminal W/L ratio Terminal leaflet shape of base Leaf color SPAD value Leaf area
(mm2)		 Petiole length
(mm)
GWEBS-05 65.7 64.5 1.02 acute 45.6 66.9 126.7
GWEBS-08 72.3 72.1 1 obtuse 39.9 109.4 147.6
GWEBS-12 75.9 75.6 1 obtuse 43.5 119.8 86.5
GWEBS-18 53.7 53 1.01 obtuse 38 70.5 99
GWEBS-29 66.9 73.9 0.91 obtuse 38.8 72.7 117.7
GWEBS-43 57.2 58.3 0.98 obtuse 29.9 84.9 156.2
GWEBS-61 70.2 69.9 1 obtuse 40.5 110.9 97.3
GWEBS-66 64 73.5 0.87 rounded 38.8 68.3 95.2
GWEBS-69 85.7 87.1 0.98 acute 37.2 68.9 125.6
GWEBS-82 61.8 58 1.07 rounded 38.1 97.7 122.4
GWEBS-90 67.6 55.1 1.23 acute 37.3 106.4 130.6
GWEBS-99 70.6 51.4 1.37 obtuse 33.9 58.7 89.4
GWEBS-111 74.1 77.5 0.96 obtuse 36.4 107.5 108.3
GWEBS-116 75.6 64.2 1.18 obtuse 37.1 72.3 139.2
GWEBS-120 64.8 59.4 1.09 obtuse 39.9 59.7 114.7
GWEBS-127 71.7 67.8 1.06 acute 38.7 63 132.6
GWEBS-128 67.8 60.4 1.12 obtuse 40.3 79 103.4
GWEBS-130 66.2 63.5 1.04 obtuse 39.8 86.2 106.6
GWEBS-147 52 58.6 0.89 obtuse 42.8 65.4 113.2
GWEBS-152 62.1 62 1 obtuse 45 109.3 93.4
Charlotte 73.7 74.4 0.99 obtuse 43.5 107.7 142.3
Flamenco 81.8 70.2 1.17 obtuse 40.4 74.7 158.2

Flower characteristics of ever-bearing strawberry crossed lines.

Lines Calyx diameter (mm) Sepal length (mm) Sepal width (mm) Sepal W/L ratio Relation between flower size and calyx Petal length (mm) Petal width (mm) Petal L/W ratio No. of petal No. of flower Flower diameter (mm) Peduncle length (cm) Flower arrangement of petal
GWEBS-05 9.2 13.2 8.4 1.56 large 8.9 8.6 1.04 6.2 8.3 19 17.1 Overlapping
GWEBS-08 9.4 9.9 5.9 1.67 same 9.1 10.6 0.86 6 9.7 24.4 14.8 Touching
GWEBS-12 12.7 12.7 7.1 1.79 large 10 12.4 0.81 6 9.7 21.3 16.6 Overlapping
GWEBS-18 12.2 9.5 5.4 1.76 large 6.8 8 0.86 5.4 10 22.5 13.4 Touching
GWEBS-29 9.8 10.2 5.6 1.82 large 8 9.3 0.86 5.4 8.7 19.6 18.7 Overlapping
GWEBS-43 11.4 8.8 5.3 1.65 large 6.3 7.6 0.83 6 10.3 13.8 16.9 Overlapping
GWEBS-61 9.7 17.4 9.1 1.9 large 8.5 9.5 0.89 5 9.7 24.8 14.6 Overlapping
GWEBS-66 12.2 17.6 11.7 1.5 large 10.6 11.3 0.93 6 10 24.5 19.2 Overlapping
GWEBS-69 11.9 14.6 8.8 1.65 large 8.2 7.4 1.11 6 9.7 26.5 20.5 Free
GWEBS-82 12.1 11.1 9.6 1.16 large 7.2 7.4 0.98 6 10 30.3 16.2 Overlapping
GWEBS-90 9.2 9 6.2 1.44 large 7.7 7.5 1.03 6 9.7 19.4 15.4 Overlapping
GWEBS-99 8.6 7.8 5.1 1.54 same 7.8 7.8 0.99 5 11.3 17.8 16.8 Overlapping
GWEBS-111 9.3 11 7 1.57 large 6.5 6.6 0.98 6 11.3 19.5 16.4 Overlapping
GWEBS-116 10.5 15.9 8.6 1.85 large 8.9 9.3 0.96 5.8 11.7 26.4 17.4 Overlapping
GWEBS-120 10.2 11.9 8.3 1.44 large 7.7 7.6 1.02 5.4 9.3 22.9 14.2 Overlapping
GWEBS-127 11 10.5 5.4 1.95 large 7.2 8 0.9 5 11.3 23.4 18.5 Overlapping
GWEBS-128 11 14.4 7.7 1.88 large 7.9 7.9 1 5.4 11.7 20.9 15.4 Overlapping
GWEBS-130 9.2 14.9 9.3 1.61 large 8.5 7.1 1.19 5 8.3 19.8 16.3 Free
GWEBS-147 8.6 7.3 4.7 1.55 large 6.6 6.3 1.04 5.4 9.7 22.6 15.8 Overlapping
GWEBS-152 9.7 8.9 5.1 1.76 same 8.6 9.5 0.91 6 11 22.4 15.7 Overlapping
Charlotte 11.7 11.7 5.7 2.05 large 9 9.4 0.96 5.2 11.7 27.5 20.2 Free
Flamenco 12 13.7 6.4 2.14 large 8.3 8 1.04 5.8 11 28.2 19.5 overlapping

Fruit characteristics of ever-bearing strawberry crossed lines.

Lines Fruit length (mm) Fruit width (mm) Fruit L/W ratio Fruit firmness (N/5 mm) Fruit weight (g) Fruit shape Fruit position of achenes Fruit position of calyx attachment Fruit attitude of sepals Sucrose Glucose Fructose citric acid Malic acid Soluble solids (°Brix)
GWEBS-05 23.7 22.4 0.95 4.21 8.6 reniform level with surface inserted outwards 8.54 5.03 19.96 7.86 0.94 7.4
GWEBS-08 31.1 29.3 0.94 5.07 7.6 ovoid level with surface level with fruit outwards 19.19 20.38 40.7 5.84 5.13 7.2
GWEBS-12 28.2 27.6 0.98 3.22 15.7 obloid level with surface inserted outwards 16.23 13.29 34.49 6.33 4.92 9.5
GWEBS-18 30.8 27.7 0.9 4.66 14.1 conical above surface inserted outwards 6.57 9.85 38.73 4.74 6.36 7.5
GWEBS-29 27.7 27.1 0.98 5.73 9.4 conical level with surface level with fruit upwards 14.4 15.02 35.87 6.75 1.52 8.2
GWEBS-43 19.7 18.7 0.95 5.03 10.8 conical above surface level with fruit upwards 7.76 7.53 25.14 6.11 0.12 6.8
GWEBS-61 29.8 26.8 0.9 3.41 8.9 ovoid above surface raised outwards 13.91 17.01 76.7 6.61 3.02 10.6
GWEBS-66 28.1 22.2 0.79 3.92 12.5 globose above surface level with fruit outwards 22.25 24.73 50.84 6.07 0.93 8.8
GWEBS-69 24.9 24.7 0.99 3.29 10.4 globose level with surface level with fruit upwards 1.08 1.25 21.24 6.45 4.03 6.5
GWEBS-82 25 21.7 0.87 3.02 11.8 conical level with surface level with fruit upwards 20.01 14.01 37.18 6.79 1.04 8.6
GWEBS-90 25.3 24.6 0.97 4.91 8.6 conical level with surface level with fruit upwards 15.72 11.09 31.14 9.2 2.73 8.8
GWEBS-99 20.2 19.7 0.98 6.01 7.9 conical below surface level with fruit upwards 5.16 3.08 11.65 5.35 3.52 6.7
GWEBS-111 19.8 18.7 0.94 5.24 7 globose level with surface level with fruit outwards 21.97 11.16 35.03 6.76 1.82 8.9
GWEBS-116 25.4 27.1 1.07 3.75 10.7 wedged level with surface level with fruit outwards 9.95 8.05 28.31 8.4 2.82 7.4
GWEBS-120 26.5 24.9 0.94 3.31 10.8 reniform below surface level with fruit outwards 2.11 2.74 21.6 5.48 1.62 8.2
GWEBS-127 32.2 29.4 0.91 5.59 12.4 conical above surface level with fruit outwards 6.55 2.16 15.33 6.27 4.37 8.9
GWEBS-128 30.7 28.3 0.92 3.75 11.4 conical level with surface level with fruit upwards 4.91 13.28 36.46 4.02 0.93 8.7
GWEBS-130 33 30.1 0.91 3.74 7.9 cordate level with surface level with fruit upwards 23.72 16.74 32.87 6.27 4.28 10
GWEBS-147 25 21.5 0.86 4.69 6.7 ovoid level with surface level with fruit outwards 27.29 19.97 46.18 7.24 3.05 7.6
GWEBS-152 25.5 25.1 0.98 4.64 9.3 conical level with surface level with fruit outwards 8.96 7.02 21.5 5.7 0.02 6.9
Charlotte 31 32.4 1.05 4.3 10.7 conical level with surface level with fruit upwards 8.48 19.74 44.67 5.02 0.83 8.2
Flamenco 34.7 31.8 0.92 4.02 11.2 conical level with surface level with fruit upwards 18.77 17.05 36.62 5.71 5.42 7.7
Table 1 Plant characteristics of ever-bearing strawberry crossed lines.
Table 2 Foliage characteristics of ever-bearing strawberry crossed lines.
Table 3 Flower characteristics of ever-bearing strawberry crossed lines.
Table 4 Fruit characteristics of ever-bearing strawberry crossed lines.

Table 1

Table 2

Table 3

Table 4

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