Strawberry Germplasm Influences Fruit Physicochemical Composition More than Harvest Date or Location

Haynes, Brianna; Fernandez, Gina; Ma, Guoying; Chen, Hsuan; Perkins-Veazie, Penelope

doi:10.3390/horticulturae11010055

Open AccessFeature PaperArticle

Strawberry Germplasm Influences Fruit Physicochemical Composition More than Harvest Date or Location

by

Brianna Haynes

^1,2,

Gina Fernandez

¹,

Guoying Ma

²

,

Hsuan Chen

¹

and

Penelope Perkins-Veazie

^1,2,*

¹

Department of Horticultural Science, North Carolina State University, Raleigh, NC 27695, USA

²

Plants for Human Health Institute, North Carolina State University, Kannapolis, NC 28081, USA

^*

Author to whom correspondence should be addressed.

Horticulturae 2025, 11(1), 55; https://doi.org/10.3390/horticulturae11010055

Submission received: 4 December 2024 / Revised: 30 December 2024 / Accepted: 3 January 2025 / Published: 7 January 2025

(This article belongs to the Special Issue Molecular Mechanisms of Fruit Quality Development and Regulation)

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Abstract

:

Strawberries (Fragaria × ananassa) are highly valued for their colorful fruit and flavorful taste. Anthocyanins provide much of the red fruit color, and the flavor is highly affected by soluble sugars and non-volatile organic acids. These fruit composition parameters impact consumer decisions. In this study, strawberry fruits from 17 commercial cultivars and advanced selections were collected weekly from replicated trials at three locations in North Carolina. The relative effects of the location and germplasm on fruit composition were determined, including the soluble solid concentration (SSC), titratable acidity (Tacid), and total anthocyanin content (TAC). The breeding criteria of at least 8.0% SSC and 0.80% Tacid were met by eight and six genotypes, respectively and five of these met both criteria. The fruit TAC ranged from 24.0 to 45.7 mg pelargonidin-3-O-glucoside (P3G) equivalents/100 g FWT. P3G was the dominant pigment in all genotypes, followed by pelargonidin-3-O-rutinoside (P3R). As harvest dates advanced, pH, TAC, P3G, P3R, and pelargonidin-3-O-(6″-malonylglucoside) (P3MG) generally decreased, while Tacid, SSC, and cyanidin-3-O-glucoside (C3G) increased. Composition of field-grown strawberries in this mid-Atlantic location were more influenced by the genotype and harvest date than by the growing location.

Keywords:

anthocyanin; cyanidin; Fragaria × ananassa; pelargonidin; quality; soluble solids; titratable acidity

Graphical Abstract

1. Introduction

The modern strawberry (Fragaria × ananassa) provides high amounts of dietary vitamin C (60–70 mg/100 g) and is valued for its sweet and distinctive flavor. Florida and California provide most of the winter and summer supply in the United States, while the South Atlantic Region (Alabama, Georgia, North Carolina, South Carolina, and Virginia) supplies spring-grown fruit [1]. North Carolina leads this region in overall production and had 485.6 hectares valued at over $26 million in 2017 [1]. North Carolina temperatures range from 0 to 38 °C during transplant establishment and fruiting, and the average rainfall is 112 cm. Therefore, North Carolina’s breeding program goals include disease-resistant germplasm, moderate chilling hours, a consistent fruit size, full color, an excellent flavor, and a good postharvest life.

The phytochemical and physicochemical properties of the fruit, along with agronomic traits, contribute to the success of commercial strawberry cultivars. Initial consumer impression of fruit quality is based on the fruit’s appearance (fruit color, shape, and size), with repeat purchases dependent on the sweetness and overall flavor [2]. The concentration of anthocyanins in the flesh and peel of strawberry fruit is correlated with the color of the fruit [3]. These water-soluble anthocyanin pigments, providing most of the visual color of strawberries, range from 10 to >40 mg pelargonidin-3-O-glucoside (P3G) equiv/100 g fresh weight (FWT) in strawberries. Most of the total anthocyanin (TAC) in strawberries are from the orange-red P3G, followed by pelargonidin-3-O-rutinoside (P3R), cyanidin-3-O-glucoside (C3G), and pelargonidin-3-O-(6″-malonylglucoside) (P3MG) [4,5,6]. The anthocyanin content can be affected by germplasm, the relative stage of ripeness, and the growing environment.

Strawberry flavor is a complex system, affected by the amount and type of sugars, amino acids, organic acids, and aroma volatiles [7,8,9]. An estimation of the soluble sugar concentration (SSC) and non-volatile organic acids or titratable acidity (Tacid) is often used as a rapid means of screening strawberry genotypes for flavor. In earlier studies, the target goals for strawberry flavor values were set at a minimum of 7.0% SSC and a maximum of 0.8% Tacid [10,11]. The current recommendations are to increase the strawberry SSC to 8.0–9.0% [7].

The range of adaptation of a strawberry selection across locations can strongly influence industry use. The preharvest environmental conditions in a production location, such as soil type, air/soil temperature, rainfall, humidity, pest pressure, and amount and quality of light, can greatly affect the physicochemical attributes of strawberries [12,13,14]. Three Korean commercial strawberry cultivars grown and harvested in three production fields were found to significantly differ in their SSC, Tacid, pH, and TAC [15]. A decreased strawberry SSC was attributed to an increased crop load rather than temperature or rainfall, while the fruits’ Tacid increased with rising temperatures [12]. Fruit redness increased with higher temperatures and decreased with rainfall [12], and the anthocyanin, phenolic, and total antioxidant content also increased with higher day/night temperatures (25–30/22 °C) [16].

The effects of harvest date and genetics on strawberry fruit composition have also been studied. Fruit development, sensory qualities, and marketability can be highly variable with harvest date, while genetics ultimately affect the response to both location and harvest date. Zhang et al. [17] reported that harvest date was a major source of variation in strawberry color, SSC, and Tacid, while variance in TAC was attributed to genetics in Asian commercial cultivars. Jouquand et al. [8] determined that fruit SSC in genotypes decreased from 9.2 to 6.4% with harvest date, while fruit Tacid showed no consistent pattern. Genetic variation appeared to outweigh harvest date, year, and location effects for fruit color, SSC, pH, Tacid, and TAC in both California [18] and Florida studies [19,20]. In a study with strawberries grown in North Carolina, ‘Liz’ (previously NCS 10-038) and ‘Rocco’ (previously NCS 10-156) fruits were slightly higher in Tacid than fruits from ‘Camarosa’ and ‘Chandler’, while SSC was highest in ‘Rocco’ fruits and TAC was lowest in ‘Liz’ fruits [21].

The individual anthocyanin pigments of strawberries may be affected by both genotype and location. Cocco et al. [22] concluded that genotype had a greater effect on the amounts of P3G and C3G pigments than production site, while Crespo et al. [23] determined that C3G, P3R, and another minor pelargonidin derivative (presumed to be P3MG) were more affected by production site than genotype. The anthocyanin profiles of ‘Liz’ and ‘Rocco’ fruits showed less %P3G and more %P3MG than ‘Camarosa’ and ‘Chandler’ fruits when all four cultivars were grown in one site [21].

As the complexities of genetics and environmental conditions of location and harvest can affect strawberry fruit composition, North Carolina germplasm grown at multiple locations and harvested over a fruiting season was examined to determine the relative genotype response. The composition of fruits from advanced selections was also compared to that of standard cultivars used in North Carolina.

2. Materials and Methods

2.1. Experimental Design and Plant Material

Seventeen strawberry genotypes (encompassing commercial cultivars and advanced selections) were used in this study. All material was grown at three locations in North Carolina over one growing season. These locations included the Central Crops Research Station (CCRS) in Clayton, NC (35.55839° N, 78.50631° W); the Piedmont Research Station (PRS) in Salisbury, NC (35.69501° N, −80.62939° W); and the North Carolina State University Horticultural Crops Research Station (HCRS) in Castle Hayne, NC (34.32051° N, −77.91533° W) (Table A1). A completely randomized block design was used at each location, with three plots per genotype and 20 plants per plot established in an annual hill plasticulture system. Plugs were planted into raised beds covered with black plastic in August–September of 2021. Preplant, fertigation, and chemical protocols followed commercially recommended practices, as outlined in the Southeast Regional Strawberry Plasticulture Production Guide [24]. Additional information on the location soil type, plant maintenance practices, and temperature/rainfall can be found in Haynes [25].

2.2. Sample Collection

Marketable fruit was collected from field trials from 25 April 2022 to 12 May 2022 (Table A1). Three harvests of five fruits per plot and per harvest were collected from each genotype at all locations as available. Marketable fruit included those that were at least 10 g in weight, free from visible defects such as disease, sunscald, or water damage, and not misshapen. Fruit were placed into plastic lock bags, frozen at −15 °C at each respective location, transported to the Plants for Human Health Institute in Kannapolis, NC, and held at −20 °C until analysis.

2.3. Fruit Compositional Analysis

All fruit used were red in the pith as well as in the epidermis. To determine whether juice or puree should be used, preliminary tests were conducted on fully thawed fruit to compare compositional values. As no differences were found between juice and puree values, juice was selected for testing strawberry genotypes. The fruit samples were thawed to room temperature and juice was collected to determine fruit composition, including the soluble solid content (SSC), pH, titratable acidity (Tacid), and anthocyanin content (TAC). A 0.5 mL aliquot of strawberry juice was placed on a handheld digital refractometer (Atago PAL-1, Bellevue, WA, USA) to determine %SSC. The pH of strawberry juice was determined using a pH meter (Thermo Scientific™ Orion Star™ A211, Waltham, MA, USA) and electrode (Thermo Scientific™ Orion™ Ross, Waltham, MA, USA). For titratable acidity, 0.5 mL of juice was diluted with 24.5 mL of distilled deionized water and thoroughly mixed, and an aliquot was placed on a digital acidity meter (Atago PAL-BX/ACID F5, Bellevue, WA, USA), with measurements expressed as % citric acid equivalents.

2.4. Anthocyanin Analysis

Samples for anthocyanin analysis were prepared using 0.4 mL of juice and 1 mL of the anthocyanin extract solvent, which consisted of UPLC-grade methanol (Fisher Scientific, Fair Lawn, NJ, USA) acidified with formic acid (Sigma-Aldrich, Burlington, MA, USA) and diluted with distilled deionized water at a ratio of 60:3:37, following the method of Perkins-Veazie et al. [21] with slight modifications. The mixture of the juice and the extract solvent was vortexed and centrifuged at 10,600× g for 20 min at 4 °C. The supernatants were filtered through 0.2 µm PTFE filters (VWR International, Radnor, PA, USA) into amber vials (Fisher Scientific, Rockwood, TN, USA) packed with N₂, sealed with screw caps (Fisher Scientific, Rockwood, TN, USA) and held at −80 °C until analyzed.

Anthocyanin separation was performed using a Waters™ ACQUITY UPLC System (Waters, Milford, MA, USA) equipped with a photodiode array (PDA) detector, a sample manager (10 °C), a column manager (45 °C), and a binary solvent manager. Empower 3 chromatography software (Waters, Milford, MA, USA) was used as the system-run controller and for data processing. Approximately 2 µL of the sample was analyzed using a reversed-phase C18 column (ACQUITY UPLC BEH C18 um, 2.1 × 100 um, Waters, Milford, MA, USA). The mobile phase consisted of 100% methanol (A1) and 5% formic acid in water (B1), with a flow rate of 0.3 mL/min using a gradient of 0 min, 100% B1; 7 min, 88% B1; 10 min, 84% B1; 15 min, 75% B1; 18 min, 60% B1; and 20 min, 100% B1.

The anthocyanin concentrations were determined using external standards of pelargonidin-3-O-glucoside (P3G) (Chromadex, Irvine, CA, USA). Standard curves were generated by injecting 1 μL of 0.00625–0.1 mg/mL preparations of the standard. Pelargonidin-3-O-rutinoside (P3R), cyanidin-3-O-glucoside (C3G), and pelargonidin-3-O-(6″-malonylglucoside) (P3MG) were identified based on previously published reports [4,6,26]. Sample anthocyanin content was reported as mg pelargonidin-3-O-glucoside/100 g fresh weight (FWT), and total anthocyanins were the sum of the identified anthocyanins. The relative content of individual anthocyanin pigments [100% × ({mg/100 g individual pigment}/{mg/100 g total anthocyanin content})] was also measured (Figure 1).

2.5. Statistical Analysis

The compositional analysis of the strawberry samples was designed as a multi-factor, completely randomized block design with three replicates (field plots) per genotype across three locations and harvest dates. The statistical analyses and visualizations of the fruit composition were performed using R (version 4.2.2, Vienna, Austria) and RStudio (version 2022.12.0+353, Boston, MA, USA). As the strawberry cultivars and selections did not yield marketable berries simultaneously, this resulted in unbalanced data. To ensure the homogeneity and normality of sample population distributions, Levene’s test for the quality of variances and Shapiro–Wilk’s test for normality were conducted for all fruit composition traits. Traits demonstrating homogeneity and normality were analyzed by analysis of variance (ANOVA), and significant differences were detected using Tukey’s honest significant difference (HSD) post hoc analysis. Nonhomogeneous effects were analyzed using Welch’s ANOVA and Games–Howell post hoc analysis, while non-normally distributed effects were analyzed using Kruskal–Wallis nonparametric ANOVA and Games–Howell post hoc analysis.

3. Results and Discussion

3.1. Strawberry Fruit Physicochemical Composition

The independent variables of genotype, harvest date, and location yielded different responses depending on the parameter measured (Table 1 and Table A2). The fruit pH and soluble solid content (SSC) values were similar across locations (Table 1). The titratable acidity (Tacid), SSC, and pH were affected uniformly by harvest date and genotype, as indicated by the lack of a significant interaction of genotype × harvest date for these parameters. However, location affected genotype response, and harvest dates did not yield similar responses across locations.

Strawberry values for pH, Tacid, and SSC were in the recommended range for ripe strawberry fruit [10], with no strong correlations found among the variables. Harvest date and genotype had the most profound effects on the physicochemical variables. A decrease in fruit pH (3.64 to 3.49) occurred for later harvest dates, with fruit from all genotypes lowest in pH from the third (last) harvest. Strawberry juice pH values were as high as 3.64–3.77 for NC20-002, NC20-008, and NC20-054 and were lowest (3.46–3.52) for ‘Liz’, NC19-020, NC20-099, and ‘Chandler’, respectively (Figure 2A). Juice pH was lowest for eleven genotypes when harvested from the Central Crops Research Station (CCRS) and highest for eight genotypes when harvested from the Piedmont Research Station (PRS).

Perceived strawberry flavor and sweetness is largely determined by the balance of Tacid and SSC [7,27]. In this study, strawberry Tacid content varied with genotype, location, and harvest date (Table 1), ranging from 0.47 to 1.17%. Six genotypes had a relatively low Tacid (0.64–0.74%) and six genotypes had average Tacid levels of at least 0.80% (Figure 2B). Fruit harvested from the Horticultural Crops Research Station (HCRS) was often highest in Tacid (0.81%), with the PRS fruit usually the lowest (0.71%). Fruit from the third harvest had the highest Tacid values across genotypes, with the exception of NC19-018, NC20-002, and ‘Ruby June’, which had the highest Tacid in the first harvest.

Strawberry SSC varied from 4.8 to 13.5%, with 15 of the 17 North Carolina genotypes having an average SSC ≥ 7.0% and eight genotypes with an SSC ≥ 8.0% across locations and harvest dates (Figure 2C). ‘Rocco’ fruit was highest in SSC at 9.3%. Generally, a higher SSC was associated with later harvest dates, with averages increasing from 7.3 to 8.6% between the first and third harvests for most of the germplasm. Strawberry SSC of North Carolina selection NC19-018 was higher (7.5–7.6%) at the HCRS and PRS locations, but lower (6.7%) at the CCRS [25]. Of the North Carolina germplasm, five genotypes (‘Chandler’, NC19-022, NC20-058, ‘Rocco’, and ‘Ruby June’) met the suggested targets of a minimum of 0.80% Tacid and 8.0% SSC [7]. Fruit from ‘Liz’, NC20-008, and NC20-055 had an SSC ≥ 8.0%, but a lower Tacid (0.68–0.77%), and fruit from NC19-018 had a Tacid of 0.82%, but a low SSC (7.2%).

The physicochemical values of the germplasm in this study were comparable to those previously reported (SSC, Tacid) for commercial cultivars grown in California [10,12], Florida [28,29], Alabama [30], and North Carolina [21]. These results indicate that the advanced selections in the North Carolina breeding program are well within the range of fruit composition values desired for the U.S. fresh market industry.

3.2. Strawberry Fruit Anthocyanin Content and Composition

A diverse range of anthocyanin values was found across genotypes (14.77–74.17 mg pelargonidin-3-O-glucoside (P3G) equivalents/100 g FWT) in this study (Table A3 and Table A4). Total anthocyanin content (TAC) is defined here as the sum of the individual anthocyanins identified by UPLC in strawberry germplasm and was most affected by genotype (Table 2). Fruit from eleven genotypes had TAC ≥ 30.0 mg/100 g, with five visually dark-colored genotypes (‘Camarosa’, NC19-016, NC19-018, NC19-020, and NC20-099) exceeding 40.0 mg/100 g (Figure 3 and Figure 4A). Those lowest in TAC were NC19-022, NC20-002, NC20-058, and ‘Ruby June’ (Figure 4A). Total anthocyanin content also varied across locations and harvest dates, with fruit from PRS highest in TAC in 15 out of 17 genotypes and TAC highest in fruit from the second harvest and lowest in the third (last) harvest.

The TAC of strawberries from North Carolina germplasm was generally higher than that reported for fruit in other studies. Strawberry fruits from commercial cultivars originating from California and Florida breeding programs have reported TAC values of 9.2–59.8 and 8.5–54.0 mg P3G/100, respectively [18,20,26,31]. ‘Camarosa’ and ‘Chandler’ are widely grown in North Carolina and are often used as standards in strawberry breeding programs. Anthocyanin values of 23.0 to 71.5–84.0 mg P3G/100 g have been reported in these genotypes [21,30,32,33]. The ‘Camarosa’ and ‘Chandler’ fruit TAC values from our study were similar to reports for germplasm from other countries [26,32,34].

As noted in previous studies for red-fruited strawberries [4,6,21], pelargonidin-3-O-glucoside (P3G) is the dominant anthocyanin pigment, representing as much as 91% of the total anthocyanin content. In our study, total P3G was heavily influenced by genetics and location (Table 2). However, fruits high in total P3G content were not always the highest in relative P3G (as %P3G of total anthocyanins). For instance, fruit from ‘Camarosa’, NC19-016, and NC19-020 had high total amounts of P3G (31.5–38.5 mg/100 g), while the highest %P3G (86.4–87.9%) was found in NC19-018, NC20-002, and NC20-054 (Figure 4B).

Other anthocyanins, including pelargonidin-3-O-rutinoside (P3R), pelargonidin-3-O-(6″-malonyl)-glucoside (P3MG), and cyanidin-3-O-glucoside (C3G), also varied significantly among genotypes (Table 2). In the North Carolina genotypes, P3G and P3R were highly correlated, and genotypes high in TAC were often high in both P3G and P3R (Figure 4A–C). The average P3R decreased from 3.5 to 2.8 mg/100 g between the first and third harvest dates, while the values were similar between the first and second harvest dates. Additionally, fruit of 10 genotypes harvested from PRS were lower in P3R, and 16 genotypes were lower in relative values than fruit from CCRS and HCRS (Table A2 and Table A3).

Several North Carolina genotypes were higher than others in the minor pigments P3MG and C3G. In this study, the P3MG content was highly dependent on the genotype, ranging from either very low (0.1–1.6%) or high (12.8–23.5%), with few values in between (Figure 4D, Table A4). Interestingly, neither the effects of location nor harvest date alone, nor their interaction together (location × harvest date), had a notable impact on this pigment (Table 2). With the exception of ‘Ruby June’, all genotypes in this study that were high in P3MG (‘Liz’, NC19-022, NC20-055, NC20-099, and ‘Rocco’) were bred in the North Carolina program. The pedigrees for ‘Rocco’ and ‘Liz’, the latest commercial releases from the North Carolina program, are available [35]. Additionally, there was no discernible correlation with TAC [36], suggesting that the presence or absence of P3MG in strawberries may be dependent on a single gene. A locus on linkage group LG6b has been hypothesized to contain a gene associated with P3MG biosynthesis in cultivated strawberry fruits, which could be responsible for the absence of P3MG in some material [5]. The C3G content varied primarily with genotype and harvest date (Table 2, Figure 4E), with the lowest amounts found in fruit from the second harvest and the highest amounts in fruit from the third harvest for 11 genotypes. Although C3G was not significantly different among locations, the significant interaction of location × harvest date indicates a differing response of harvest date with location (Table 2).

The possible differences in TAC and in anthocyanin profiles among strawberry studies may be influenced by fruit ripeness definition and selection. Strawberries used for fresh market must have a minimum color of three-quarters red [8,17,18,19]. However, in North Carolina, direct market and local sales outlets prefer fully red fruit [1], and fully red fruit were used in our study. Total anthocyanin content can increase by as much as 31% as fruit ripen from ¾ to full color [33].

3.3. Effects of Genotype, Harvest Date, and Location

The strawberry fruit composition components were affected nonuniformly by location, harvest, and genotype factors. Overall, genotype (collectively encompassing commercial cultivars and advanced selections) had the greatest effect on fruit composition, followed by harvest date, similar to results of Pelayo-Zaldívar et al., Cayo et al., and Kelly et al. [18,19,20]. Several QTLs associated with strawberry fruit total and individual sugars, soluble solid content, organic acids, titratable acidity, anthocyanins, pH, and other fruit quality measurements have been identified [37,38,39,40]. These QTLs support our findings by providing a genetic basis for variation in strawberry fruit composition parameters, further emphasizing that genetics are primarily responsible for differences observed in strawberry fruit composition. The germplasm had a greater and more consistent influence across harvest dates and locations, as indicted by significant genotype effects for most measured traits.

In our study, Tacid, SSC, C3G, and %C3G content generally increased with later harvest dates, while pH, TAC, P3G, P3R, P3MG, %P3G, %P3R, and %P3MG decreased. Notable differences were found for SSC (7.3, 7.7, and 8.6%) and Tacid (0.76, 0.73, and 0.79%) for harvests 1, 2, and 3, respectively [25]. The fruit composition values averaged across genotypes and locations were most often similar between the first and second harvest dates, while the third date statistically differed from one or both previous dates. The location had the least influence on strawberry fruit composition, and was significant for Tacid, TAC, P3G, and P3R. Among the three locations, fruit from PRS was most often different than that from CCRS or HCRS stations, with differences more pronounced during specific harvest dates. Only the Tacid values statistically differed at all three research stations. Differences in the other fruit composition variables could not be statistically attributed to location alone. Additional genotype, harvest date, and location effects on strawberry fruit composition are detailed in Table A2, Table A3 and Table A4.

The two-way interaction between genotype × harvest date was not significant, indicating a consistent impact of harvest date across genotypes. This is supported by the fact that fruit from all the genotypes was collected on the same date at each location, with the three harvests occurring within an 11- to 17-day timeframe due to the small fruiting window in North Carolina. However, significant location × harvest date and location × genotype interactions for all physicochemical and most phytochemical variables measured illustrate the need to test advanced selections in multiple locations and for multiple harvest dates to develop a comprehensive profile.

4. Conclusions

The strawberry fruit composition of twelve advanced selections from the North Carolina strawberry breeding program and five commercial genotypes were found to vary primarily with genotype, followed by harvest date. Five genotypes/advanced selections met the desired % soluble solid content (SSC) (≥8.0) and % titratable acidity (Tacid) (0.80) target values. Pelargonidin-3-O-glucoside (P3G) was the dominant anthocyanin pigment (60.2 to 90.7%) in all the genotypes, followed by pelargonidin-3-O-rutinoside (P3R) (3.5 to 17.4%). Several North Carolina genotypes were unusually high in the minor pigment pelargonidin-3-O-(6″-malonylglucoside) (P3MG). Our results indicate that the genotype and harvest date, rather than the production location, most affect strawberry fruit composition in North Carolina.

Author Contributions

Conceptualization, B.H., G.F., and P.P.-V.; methodology, G.F., P.P.-V., G.M. and B.H.; software, B.H. and G.F.; validation, B.H. and H.C.; formal analysis, B.H. and H.C.; investigation, B.H.; resources, G.F., P.P.-V. and H.C.; data curation, B.H.; writing—original draft preparation, B.H.; writing—review and editing, B.H., G.F., G.M., H.C. and P.P.-V.; visualization, B.H. and H.C.; supervision, G.F. and P.P.-V.; project administration, P.P.-V.; funding acquisition, P.P.-V. and G.F. All authors have read and agreed to the published version of the manuscript.

Funding

This research was supported by USDA NIFA Hatch #7005040 and the North Carolina Strawberry Association 15-01-57002.

Data Availability Statement

The original data presented in this study are openly available at the North Carolina State University library, theses and dissertations, URI: https://www.lib.ncsu.edu/resolver/1840.20/41885, accessed on 2 January 2025.

Acknowledgments

We thank Joyce Edwards, Erin Deaton, Sheridan Moore, and Rocco Schiavone for their technical help and the support personnel at the Piedmont Research Station, the Central Crops Research Station, and the North Carolina State University Horticultural Crops Research Station.

Conflicts of Interest

The authors declare no conflicts of interest.

Appendix A

Table A1. Locations of replicated trials where fruit samples of each genotype were harvested for the 2021–2022 season in North Carolina.

Station	Longitude	Latitude	City	Soil Type	Planting	Harvest
CCRS ¹	35.66839°	−78.50631°	Clayton	Wagram loamy sand (0–6%) ²	11 October 2021	25 April 2022
						5 May 2022
						12 May 2022

HCRS ¹	34.32051°	−77.91533°	Castle Hayne	Torhunta loamy fine sand (0–2%)	26 October 2021	28 April 2022
				Sallings fine sand (2%)		5 May 2022
						9 May 2022

PRS ¹	35.69501°	−80.62939°	Salisbury	Llyod clay loam (2–8%)	1 September 2021	25 April 2022
						2 May 2022
						12 May 2022

¹ CCRS = Central Crops Research Station, Clayton, NC; HCRS = Horticultural Crops Research Station, Castle Hayne, NC; PRS = Piedmont Research Station, Salisbury, NC. ² Soil type percentages indicate the slope of the field.

Table A2. Mean values for fruit composition variables of 17 commercial strawberry genotypes and advanced selections grown at three locations in North Carolina (2022).

		pH	Tacid (% as Citric Acid) ¹	SSC (%) ¹
Source	Categories		Mean ± SD
Location ²	CCRS ¹	3.58 ± 0.15 a	0.77 ± 0.13 b	7.9 ± 1.29 a
	HCRS ¹	3.57 ± 0.11 a	0.81 ± 0.09 a	7.9 ± 1.14 a
	PRS ¹	3.60 ± 0.14 a	0.71 ± 0.08 c	7.8 ± 1.34 a

HDate ³	1st	3.64 ± 0.12 a	0.76 ± 0.11 b	7.3 ± 1.01 c
	2nd	3.63 ± 0.12 a	0.73 ± 0.12 c	7.7 ± 1.18 b
	3rd	3.49 ± 0.10 b	0.79 ± 0.10 a	8.6 ± 1.19 a

Genotype	Camarosa	3.60 ± 0.11 bcde	0.74 + 0.10 bcde	7.9 ± 0.89 cdef
	Chandler	3.52 ± 0.10 def	0.81 ± 0.07 abc	8.2 ± 0.79 bcde
	Liz	3.46 ± 0.10 f	0.77 ± 0.09 abcd	8.1 ± 0.98 bcde
	Miss Jo	3.63 ± 0.12 bcd	0.77 ± 0.07 abcd	7.9 ± 1.09 cdef
	NC19-016	3.55 ± 0.10 cdef	0.75 ± 0.09 bcde	6.3 ± 0.92 g
	NC19-018	3.52 ± 0.10 def	0.82 ± 0.09 ab	7.2 ± 1.05 f
	NC19-020	3.61 ± 0.16 bcde	0.64 ± 0.09 f	6.1 ± 0.84 g
	NC19-022	3.55 ± 0.12 cdef	0.81 ± 0.10 abc	8.8 ± 0.95 abc
	NC20-002	3.67 ± 0.10 ab	0.76 ± 0.09 abcde	7.8 ± 0.89 def
	NC20-008	3.77 ± 0.11 a	0.68 ± 0.12 ef	8.4 ± 1.38 abcd
	NC20-018	3.62 ± 0.11 bcd	0.71 ± 0.12 def	7.5 ± 1.09 def
	NC20-054	3.64 ± 0.13 bc	0.72 ± 0.10 cdef	7.6 ± 0.61 def
	NC20-055	3.60 ± 0.13 bcde	0.74 ± 0.11 bcde	8.3 ± 1.13 bcd
	NC20-058	3.57 ± 0.09 bcde	0.85 ± 0.10 a	8.8 ± 0.94 ab
	NC20-099	3.50 ± 0.14 ef	0.75 ± 0.10 bcde	7.2 ± 0.84 f
	Rocco	3.55 ± 0.10 cdef	0.81 ± 0.09 abc	9.3 ± 1.11 a
	Ruby June	3.62 ± 0.11 bcd	0.81 ± 0.10 abc	8.2 ± 0.74 bcd

¹ CCRS = Central Crops Research Station (Clayton, NC), HCRS = Horticultural Crops Research Station (Castle Hayne, NC), PRS = Piedmont Research Station (Salisbury, NC), Tacid = titratable acidity, and SSC = soluble solids content. ² Values averaged within location. Different letters within treatment indicate statistically significantly differences at p < 0.05. ³ Values averaged within harvest date. Different letters within treatment indicate statistically significantly differences at p < 0.05.

Table A3. Mean values for total anthocyanin content and profiles of 17 commercial strawberry genotypes and advanced selections grown at three locations in North Carolina (2022).

		TAC ¹	C3G ¹	P3G ¹	P3R ¹	PM3G ¹
Source	Categories			Mean ± SD
Location ²	CCRS ¹	33.4 ± 13.0 ab	1.48 ± 1.0 a	25.97 ± 10.2 b	3.56 ± 1.9 a	2.07 ± 3.3 a
	HCRS ¹	31.13 ± 8.6 b	1.38 ± 0.9 a	24.42 ± 7.3 b	3.31 ± 1.6 a	1.89 ± 2.5 a
	PRS ¹	36.13 ± 12.8 a	1.56 ± 1.1 a	29.14 ± 11.4 a	2.90 ± 1.5 b	1.90 ± 2.8 a

HDate ³	1st	35.79 ± 12.2 a	1.49 ± 1.1 ab	28.30 ± 9.5 a	3.49 ± 1.7 a	2.15 ± 3.4 a
	2nd	33.92 ± 11.8 ab	1.38 ± 0.9 b	26.96 ± 10.5 ab	3.49 ± 1.9 a	1.77 ± 2.4 a
	3rd	31.03 ± 10.9 b	1.57 ± 1.1 a	24.42 ± 9.6 b	2.76 ± 1.3 b	1.92 ± 2.7 a
Genotype	Camarosa	40.06 ± 9.8 a	2.51 ± 0.5 b	31.53 ± 8.0 bc	5.53 ± 1.6 ab	0.21 ± 0.1 c
	Chandler	35.19 ± 11.5 abc	1.38 ± 0.9 cdef	30.46 ± 9.6 bc	3.04 ± 1.3 cdef	0.13 ± 0.1 c
	Liz	36.13 ± 9.5 abc	3.82 ± 0.8 a	24.05 ± 6.6 cdefg	3.56 ± 1.3 cde	4.64 ± 1.5 b
	Miss Jo	24.32 ± 5.4 bcd	0.46 ± 0.2 gh	19.00 ± 4.1 efg	4.24 ± 1.2 bc	0.38 ± 0.2 c
	NC19-016	45.23 ± 5.7 a	0.64 ± 0.3 fgh	37.18 ± 4.3 ab	6.49 ± 1.7 a	0.16 ± 0.1 c
	NC19-018	44.23 ± 10.3 a	1.79 ± 0.5 bcde	38.96 ± 9.6 a	2.82 ± 0.7 cdef	0.19 ± 0.1 c
	NC19-020	45.73 ± 12.6 a	2.10 ± 0.8 bc	38.52 ± 10.9 a	3.87 ± 1.4 bcd	0.13 ± 0.1 c
	NC19-022	23.02 ± 4.3 cd	0.28 ± 0.3 h	16.55 ± 3.6 fg	2.36 ± 0.6 defg	3.52 ± 0.6 b
	NC20-002	20.61 ± 4.0 d	1.93 ± 0.5 bcd	17.83 ± 3.6 efg	0.71 ± 0.2 g	0.02 ± 0.0 c
	NC20-008	33.96 ± 7.3 abc	1.76 ± 0.5 bcde	28.10 ± 6.2 cdef	3.49 ± 0.9 cde	0.32 ± 0.1 c
	NC20-018	24.48 ± 3.5 bcd	1.25 ± 0.3 defg	20.87 ± 3.4 efg	1.65 ± 0.3 fg	0.39 ± 0.1 c
	NC20-054	33.46 ± 11.2 abc	1.05 ± 0.2 efgh	29.02 ± 9.7 cd	3.13 ± 1.3 cdef	0.07 ± 0.1 c
	NC20-055	37.80 ± 5.3 ab	0.76 ± 0.3 fgh	28.49 ± 4.1 cd	3.59 ± 0.7 cde	4.91 ± 1.1 b
	NC20-058	24.00 ± 4.6 bcd	1.11 ± 0.3 defgh	20.48 ± 4.2 efg	2.13 ± 0.2 defg	0.21 ± 0.1 c
	NC20-099	40.64 ± 16.1 a	0.83 ± 0.4 fgh	27.25 ± 11.0 cdef	3.62 ± 1.6 cde	7.98 ± 5.0 a
	Rocco	37.31 ± 8.2 abc	0.83 ± 0.6 fgh	28.12 ± 7.3 cdef	3.25 ± 0.5 cdef	5.16 ± 1.0 b
	Ruby June	24.09 ± 5.4 bcd	2.58 ± 0.6 b	15.07 ± 3.6 g	2.01 ± 0.4 efg	4.11 ± 0.8 b

¹ CCRS = Central Crops Research Station (Clayton, NC), HCRS = Horticultural Crops Research Station (Castle Hayne, NC), PRS = Piedmont Research Station (Salisbury, NC), TAC = total anthocyanin content, C3G = cyanidin 3-O-glucoside, P3G = pelargonidin-3-O-glucoside, P3R = pelargonidin 3-O-rutinoside, and P3MG = pelargonidin-3-O-(6″-malonyl)-glucoside. Units in mg/100 g FWT. ² Values averaged within location. Different letters within treatment indicate statistically significantly differences at p < 0.05. ³ Values averaged within harvest date. Different letters within treatment indicate statistically significantly differences at p < 0.05.

Table A4. Mean values for anthocyanin content and individual anthocyanin profiles of 17 commercial strawberry genotypes and advanced selections grown at three locations in North Carolina (2022).

		TAC ¹	%C3G ¹	%P3G ¹	%P3R ¹	%P3MG ¹
Source	Categories			Mean ± SD
Location ²	CCRS ¹	33.4 ± 13.0 ab	4.72 ± 3.2 a	78.10 ± 7.5 b	10.49 ± 3.7 a	5.86 ± 7.4 a
	HCRS ¹	31.13 ± 8.6 b	4.79 ± 3.4 a	78.36 ± 8.3 b	10.35 ± 3.3 a	6.14 ± 7.6 a
	PRS ¹	36.13 ± 12.8 a	4.62 ± 3.6 a	79.96 ± 8.8 a	8.01 ± 3.5 b	5.27 ± 7.2 a

HDate ³	1st	35.79 ± 12.2 a	4.45 ± 3.0 b	79.29 ± 7.8 a	9.65 ± 3.5 a	5.73 ± 7.5 a
	2nd	33.92 ± 11.8 ab	4.29 ± 3.2 b	78.86 ± 8.2 a	10.18 ± 3.9 a	5.54 ± 7.0 a
	3rd	31.03 ± 10.9 b	5.38 ± 3.9 a	78.34 ± 8.7 a	8.91 ± 3.5 b	5.96 ± 7.7 a

Genotype	Camarosa	40.06 ± 9.8 a	6.44 ± 1.3 b	78.62 ± 1.5 de	13.77 + 1.6 b	0.53 ± 0.3 c
	Chandler	35.19 ± 11.5 abc	3.80 ± 1.5 cdef	86.71 ± 2.3 ab	8.70 ± 2.2 cde	0.39 ± 0.4 c
	Liz	36.13 ± 9.5 abc	11.02 ± 2.9 a	66.40 ± 2.2 h	9.62 ± 1.7 cd	12.78 ± 1.7 b
	Miss Jo	24.32 ± 5.4 bcd	1.93 ± 0.84 efg	78.19 + 2.39 ef	17.37 ± 2.19 a	1.46 ± 0.76 c
	NC19-016	45.23 ± 5.7 a	1.44 ± 0.8 fg	82.33 + 2.2 cd	14.19 ± 2.4 ab	0.36 ± 0.2 c
	NC19-018	44.23 ± 10.3 a	4.13 ± 1.1 cde	87.89 ± 2.1 a	6.45 ± 1.2 de	0.44 ± 0.4 c
	NC19-020	45.73 ± 12.6 a	4.54 ± 1.0 bcd	84.04 + 2.8 bc	8.84 ± 3.5 cde	0.25 ± 0.2 c
	NC19-022	23.02 ± 4.3 cd	1.10 ± 1.0 g	71.64 + 4.0 g	10.33 ± 2.5 c	15.33 ± 1.5 ab
	NC20-002	20.61 ± 4.0 d	9.46 ± 2.3 a	86.42 ± 2.2 ab	3.45 ± 0.6 f	0.07 ± 0.2 c
	NC20-008	33.96 ± 7.3 abc	5.20 ± 0.9 bc	82.62 ± 2.3 c	10.43 ± 2.4 c	0.98 ± 0.5 c
	NC20-018	24.48 ± 3.5 bcd	5.19 ± 1.2 bc	85.09 ± 1.7 abc	6.83 ± 1.4 de	1.60 ± 0.3 c
	NC20-054	33.46 ± 11.2 abc	3.36 ± 1.0 cdefg	86.68 ± 2.0 ab	9.39 ± 1.8 cde	0.15 ± 0.3 c
	NC20-055	37.80 ± 5.3 ab	2.03 ± 0.8 efg	75.34 ± 1.9 efg	9.58 ± 1.7 cde	12.92 ± 1.7 b
	NC20-058	24.00 ± 4.6 bcd	4.65 ± 1.3 bcd	85.26 ± 1.4 abc	9.05 ± 1.2 cde	0.83 ± 0.4 c
	NC20-099	40.64 ± 16.1 a	1.91 ± 0.9 efg	66.86 ± 1.6 h	8.80 ± 1.2 cde	17.33 ± 9.2 a
	Rocco	37.31 ± 8.2 abc	2.23 ± 1.6 defg	74.61 ± 3.2 fg	9.11 ± 2.5 cde	13.95 ± 1.5 ab
	Ruby June	24.09 ± 5.4 bcd	10.80 ± 1.6 a	62.47 ± 1.9 i	8.47 ± 1.2 cde	17.17 ± 1.3 a

¹ CCRS = Central Crops Research Station (Clayton, NC), HCRS = Horticultural Crops Research Station (Castle Hayne, NC), PRS = Piedmont Research Station (Salisbury, NC), TAC = total anthocyanin content, %C3G = cyanidin 3-O-glucoside, %P3G = pelargonidin-3-O-glucoside, %P3R = pelargonidin 3-O-rutinoside, and %P3MG = pelargonidin-3-O-(6″-malonyl)-glucoside. Percentages of anthocyanin are relative to total anthocyanin [100% × ({mg/100 g individual pigment}/{mg/100 g total anthocyanin content})]. ² Values averaged within location. Different letters within treatment indicate statistically significantly differences at p < 0.05. ³ Values averaged within harvest date. Different letters within treatment indicate statistically significantly differences at p < 0.05.

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Figure 1. UPLC chromatogram of anthocyanin extract from strawberry fruit. Peaks identified are cyanidin-3-O-glucoside (C3G), pelargonidin-3-O-glucoside (P3G), pelargonidin-3-O-rutinoside (P3R), and pelargonidin-3-O-(6″-malonylglucoside) (P3MG).

Figure 2. Mean pH (A), titratable acidity (% Tacid) (B), and soluble solids content (% SSC) (C) of all fruit samples averaged across harvest dates and locations. Dashed lines represent historical values before 2015 (7.0% SSC and maximum 0.80% Tacid) and solid lines represent current breeding aims (8.0% SSC and average 0.80% Tacid). The Tukey’s honest significant difference (HSD) results are listed at p < 0.05. Blue and red represent low and high genotypes. Error bars represent ±1 standard deviation from the mean.

Figure 3. Fruit from Piedmont Research Station (Salisbury, NC, USA) harvest, representing different scarlet, red, and dark-red colors and anthocyanin levels within North Carolina germplasm collection.

Figure 4. Mean total anthocyanin content (TAC) [mg pelargonidin-3-O-glucoside (P3G) equiv/100 g fresh weight (FWT)] (A), pelargonidin-3-O-glucoside (P3G) (mg/100 g) (B), pelargonidin-3-O-rutinoside (P3R) (mg/100 g) (C), pelargonidin-3-O-(6″-malonyl)-glucoside (P3MG) (mg/100 g) (D), and cyanidin-3-O-glucoside (C3G) (mg/100 g) (E) of all fruit samples averaged across harvest dates and locations. Tukey’s honest significant difference (HSD) results are listed at p < 0.05. Blue and red represent low and high genotypes. Error bars represent ±1 standard deviation from the mean.

Table 1. Location, harvest date, and genotype effects on fruit composition variable means of 17 commercial strawberry genotypes and advanced selections grown at three locations in North Carolina (2022).

	pH	Tacid (% as Citric Acid Equivalents) ¹	SSC (%) ¹
Source		Mean ± SD
Location	NS ²	***	NS
Harvest Date	***	***	***
Genotype	***	***	***
Location × Harvest Date	***	***	***
Location × Genotype	**	***	***
Harvest Date × Genotype	NS	NS	NS

¹ Tacid = titratable acidity and SSC = soluble solids content. ² *, **, and *** indicates statistically significant differences between factors at p < 0.05, p < 0.01, or p < 0.001, respectively. NS (not significant) indicates that the statistical difference was p > 0.05.

Table 2. Effect of location, harvest date, and genotype on mean values for total anthocyanin content and profiles of 17 strawberry commercial genotypes and advanced selections grown at three locations in North Carolina (2022).

	TAC ¹	C3G ¹	P3G ¹	P3R ¹	P3MG ¹	%C3G ²	%P3G ²	%P3R ²	%P3MG ²
Source					Mean ± SD
Location	** ³	NS	***	***	NS	NS	***	***	NS
HDate	*	*	*	***	NS	***	NS	***	NS
Genotype	***	***	***	***	***	***	***	***	***
Location × HDate	*	***	*	**	NS	**	*	***	NS
Location × Genotype	*	NS	*	*	NS	NS	*	*	NS
HDate × Genotype	NS	NS	NS	NS	NS	NS	NS	NS	NS

¹ Amounts of individual anthocyanin pigments. TAC = total anthocyanin content, C3G = cyanidin-3-O-glucoside, P3G = pelargonidin-3-O-glucoside, P3R = pelargonidin-3-O-rutinoside, P3MG = pelargonidin-3-O-(6″-malonyl)-glucoside. Units are in mg/100 g FWT. ² Relative amounts of individual anthocyanin pigments to total anthocyanin content (TAC). %C3G = cyanidin-3-O-glucoside, %P3G = pelargonidin-3-O-glucoside, %P3R = pelargonidin-3-O-rutinoside, %P3MG = pelargonidin-3-O-(6″-malonyl)-glucoside. ³ *, **, *** indicates statistically significant differences between factors at p < 0.05, p < 0.01, or p < 0.001, respectively. NS (not significant) indicates that the statistical difference was p > 0.05.

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Haynes, B.; Fernandez, G.; Ma, G.; Chen, H.; Perkins-Veazie, P. Strawberry Germplasm Influences Fruit Physicochemical Composition More than Harvest Date or Location. Horticulturae 2025, 11, 55. https://doi.org/10.3390/horticulturae11010055

AMA Style

Haynes B, Fernandez G, Ma G, Chen H, Perkins-Veazie P. Strawberry Germplasm Influences Fruit Physicochemical Composition More than Harvest Date or Location. Horticulturae. 2025; 11(1):55. https://doi.org/10.3390/horticulturae11010055

Chicago/Turabian Style

Haynes, Brianna, Gina Fernandez, Guoying Ma, Hsuan Chen, and Penelope Perkins-Veazie. 2025. "Strawberry Germplasm Influences Fruit Physicochemical Composition More than Harvest Date or Location" Horticulturae 11, no. 1: 55. https://doi.org/10.3390/horticulturae11010055

APA Style

Haynes, B., Fernandez, G., Ma, G., Chen, H., & Perkins-Veazie, P. (2025). Strawberry Germplasm Influences Fruit Physicochemical Composition More than Harvest Date or Location. Horticulturae, 11(1), 55. https://doi.org/10.3390/horticulturae11010055

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Strawberry Germplasm Influences Fruit Physicochemical Composition More than Harvest Date or Location

Abstract

1. Introduction

2. Materials and Methods

2.1. Experimental Design and Plant Material

2.2. Sample Collection

2.3. Fruit Compositional Analysis

2.4. Anthocyanin Analysis

2.5. Statistical Analysis

3. Results and Discussion

3.1. Strawberry Fruit Physicochemical Composition

3.2. Strawberry Fruit Anthocyanin Content and Composition

3.3. Effects of Genotype, Harvest Date, and Location

4. Conclusions

Author Contributions

Funding

Data Availability Statement

Acknowledgments

Conflicts of Interest

Appendix A

References

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