Research Article

Horticultural Science and Technology. 30 June 2024. 326-338
https://doi.org/10.7235/HORT.20240028

ABSTRACT


MAIN

  • Introduction

  • Materials and Methods

  •   Plant material and sample collection

  •   Analysis of volatile compounds

  •   Data analysis

  • Results and Discussion

  •   Analysis of volatile compounds by cultivar and maturity stage

  •   Analysis of volatile compounds in F. vesca

  • Conclusion

Introduction

Strawberries are one of the most popular fruits worldwide, with global production of 23.5 tons ha-1 in 2021 (FAOSTAT, 2021). They are rich in vitamin C and have a unique balance of sour and sweet flavors, along with a distinct aroma (Schieberle et al., 1997; Ulrich et al., 1997; Lee et al., 2003). The flesh of strawberries consumed by humans consists of an enlarged receptacle (Hancock, 2020). During strawberry fruit ripening, auxins promote the accumulation of fructose derivatives and secondary products (Fait et al., 2008; Vallarino et al., 2018), with sucrose, glucose, and fructose being the most abundant soluble sugars (Halford et al., 2011). Sucrose, amino acids, phenolic compounds, and volatile compounds are major indicators of fruit ripening and play an important role in determining fruit quality (Li et al., 2019; Fan et al., 2021). Furthermore, volatile compounds are basic aromatic components that not only act as attracting substances for moisture mediators but also as antibacterial agents against pathogenic fungi and bacteria (Arroyo et al., 2007; Xu et al., 2019).

Strawberries develop under the influence of auxins and ripen through three non-climacteric stages: an increase in the fruit size, an increased weight, and a color change, which occur approximately 40 days after flowering (Zhang et al., 2011; Karki et al., 2024). Strawberry fruit ripening is a complex process that involves genetic and environmental factors leading to biochemical and physiological changes in the fruit (Giovannoni, 2001; Giovannoni, 2004). Changes in color, texture, flavor, aroma, and nutrients occur during ripening, and maturation is influenced by external and internal factors (Moya-León et al., 2019). As strawberries ripen, several sugars and organic acids accumulate, and volatile compounds are released (Menager et al., 2004), significantly impacting the fruit quality, post-harvest quality, and consumer preferences (Alvarez-Suarez et al., 2014).

Flavor is a perceptual response that combines taste, smell, and touch (Prescott, 2004). The flavors of various fruits, including strawberries, are influenced by factors such as sugars, acids, fruit color, textures, and volatile compounds (Hall, 1968; Christensen, 1983). The flavor of strawberry is produced by a complex mixture of numerous volatile and functional compounds, with more than 280 volatile compounds having been identified in mature cultivated strawberries (Ulrich et al., 2018). Low-molecular-weight volatile compounds produced during ripening provide a fruity flavor, with methyl and ethyl esters accounting for 25 to 90% (Fan et al., 2021). The fruity flavor of strawberries is affected by the genotype and environment (Forney et al., 2000), and wild strawberries tend to have a stronger flavor than cultivated strawberries (Ulrich et al., 2007). The biosynthetic pathways, enzymes, and regulatory mechanisms underlying the accumulation of volatile compounds in Fragaria have been partially elucidated, and it is known that the composition of volatile compounds varies with the cultivar, developmental stage, and with certain post-harvest factors and the analytical techniques used (Urrutia et al., 2017).

In Korea, strawberries are cultivated over an area of 5,692 ha, with a production volume of 157,784 tons per year (KOSIS, 2022). Presently, the domestic cultivar penetration rate is 96.3%, with the ‘Sulhyang’ cultivar ranking first (84.5%) in terms of cultivar share, followed by ‘Keumsil’, ‘Jukhyang’, and ‘Maehyang’ (RDA, 2022). ‘Sulhyang’ is a forced culture cultivar developed by crossing ‘Yukbo’ and ‘Janghee’ at the Nonsan Strawberry Experiment Station of the Chungnam Agricultural Research and Extension Services in 2005. It is resistant to powdery mildew, juicy, and has a soft texture (Kim et al., 2006). ‘Keumsil’, bred in 2007 by crossing ‘Maehyang’, which has high flesh hardness, and ‘Sulhyang’, which offers high yields, is a cultivar suitable for forcing culture. It has an upright growth habit, elliptical leaves, strong vigorous growth, and cone-shaped, light red fruits (Yoon et al., 2020). In 2012, the Damyang Agricultural Technology Center developed a new strawberry cultivar, ‘Jukhyang’, by crossing ‘Red Pearl’ and ‘Maehyang’. It is a cultivar suitable for forcing culture due to its low dormancy, high yield, and good fruit quality (Lee et al., 2014). ‘Maehyang’ is a new strawberry cultivar released in 2002 at the ARES Nonsan Strawberry Experiment Station in South Chungcheong Province, having originated as a cross between ‘Tochinomine’ and ‘Akihime’. It has several excellent characteristics, including weak dormancy, vigorous growth, a high yield and good fruit quality, making it a suitable variety for forcing culture (Kim et al., 2004).

A number of studies have sought to analyze the variation and contents of the bioactive compounds in domestic strawberry cultivars (Kim et al., 2013; Kim et al., 2015). However, although strawberry is an important horticultural crop in Korea, studies of volatile compounds in strawberry fruit are limited. Therefore, four domestic strawberry cultivars and two European wild cultivars were used to determine differences in the qualitative characteristics of the volatile compounds in strawberries according to the maturation stage and cultivar.

Materials and Methods

Plant material and sample collection

Six cultivars of strawberry, specifically ‘Keumsil’, ‘Maehyang’, ‘Sulhyang’, ‘Arihyang’, Fragaria vesca ‘Yellow Wonder’, and F. vesca ‘Baron Solemacher’, were cultivated in this study. Strawberry seedlings were planted at 18-cm intervals in two rows in a tunnel-type greenhouse at Kyungpook National University (35°53'N 128°36'E). Seedlings of the ‘Keumsil’, ‘Maehyang’, ‘Sulhyang’, and ‘Arihyang’ cultivars were obtained from the Kyungnam Agriculture Research Station (Jinju, Korea), while seeds of F. vesca ‘Yellow Wonder’ and F. vesca ‘Baron Solemacher’ were purchased from a commercial seed company (123Seeds, Oost-Souburg, Netherlands). The experiment conducted here lasted from September 14, 2020 to March 29, 2021, with ‘Keumsil’ transplanted on September 14, 2020 and ‘Maehyang’, ‘Sulhyang’, and ‘Arihyang’ transplanted on September 21, 2020. F. vesca ‘Yellow Wonder’ and F. vesca ‘Baron Solemacher’ were sown on September 19, 2019 and April 2, 2020, respectively, and were transplanted on September 21, 2020. The soil type on which strawberries were grown was well-drained, silty soil mulched with black polyethylene film, with a base fertilizer applied at 2000 kg per 1000 m2 of commercial organic matter according to local recommendations. Irrigation was performed at regular intervals using a nutrient solution (N‑P‑K‑Ca‑Mg‑S = 16‑4‑8‑4-mEq·L-1). The experimental design used was a randomized complete block design, with five replications in the strawberry cultivars ‘Keumsil’, ‘Maehyang’, ‘Sulhyang’, and ‘Arihyang’ (20 strawberry transplants for each replication).

Fruit samples from the ‘Keumsil’, ‘Maehyang’, ‘Sulhyang’, and ‘Arihyang’ cultivars were collected at four maturity stages, as described by Wang et al. (2021), and only mature fruits were collected from the F. vesca ‘Yellow Wonder’ and F. vesca ‘Baron Solemacher’ cultivars (Fig. 1). Three fruits per cultivar, per replicate, per stage (n = 3) were sampled in the field (Table 1). Strawberry fruits in the white, green, pink, and red stages of ‘Keumsil’, ‘Maehyang’, ‘Sulhyang’, and ‘Arihyang’ were collected on February 3, 4, 5, and 9, 2021, respectively. The fruits from F. vesca ‘Yellow Wonder’ and F. vesca ‘Baron Solemacher’ were collected on February 8, 2021. All collected samples were frozen at ‒50 degrees C using a freezer and then analyzed.

https://static.apub.kr/journalsite/sites/kshs/2024-042-03/N0130420309/images/HST_42_03_09_F1.jpg
Fig. 1.

Green, white, pink, and red stages of the ‘Sulhyang’ (A), ‘Maehyang’ (B), ‘Keumsil’ (C), ‘Arihyang’ (D) and Fragaria vesca ‘Yellow Wonder’ and F. vesca ‘Baron Solemacher’ (E) cultivars.

Table 1.

Sampling stages, dates, replicates, and number of fruits for strawberry cultivars

Cultivar Sampling stage Sampling date Replicates (fruit)
‘Keumsil’ White 3 Feb. 2021 3
Green 4 Feb. 2021 3
Pink 5 Feb. 2021 3
Red 9 Feb. 2021 3
‘Maehyang’ White 3 Feb. 2021 3
Green 4 Feb. 2021 3
Pink 5 Feb. 2021 3
Red 9 Feb. 2021 3
‘Sulhyang’ White 3 Feb. 2021 3
Green 4 Feb. 2021 3
Pink 5 Feb. 2021 3
Red 9 Feb. 2021 3
‘Arihyang’ White 3 Feb. 2021 3
Green 4 Feb. 2021 3
Pink 5 Feb. 2021 3
Red 9 Feb. 2021 3
F. vesca ‘Yellow Wonder’ - 8 Feb. 2021 3
F. vesca ‘Baron Solemacher’ - 8 Feb. 2021 3

Analysis of volatile compounds

The analysis of the volatile compounds was performed as previously described (Rambla et al., 2017). Approximately 3 g of frozen fruit in each case was sampled using a cork borer, with the sample placed in a 10 mL headspace glass vial and incubated in a 37°C water bath for 10 min. Then, 6.6 g of CaCl2·2H2O and 3 mL of 100 mM EDTA (pH 7.5) were added and sonicated for 5 min. Samples in the headspace glass vials were preheated at 50°C for 10 min. A 65 µm polydimethylsiloxane–divinylbenzene fiber (Supelco, Inc., Bellefonte, PA, USA) was exposed to the vial headspace at 50°C for 10 min to extract volatile compounds. The volatile compound analysis was conducted using a gas chromatograph–mass spectrometer (7890B-5977B GC/MSD, Agilent, Santa Clara, CA, USA) at the Instrumental Analysis Center of Kyungpook National University (Daegu, Korea). This analysis was performed within 12 hours of extraction. Volatile compounds were desorbed from the injection port of the gas chromatograph at 250°C for one minute in the splitless mode. Separations were performed on a DB-5 ms column (60 m × 0.25 mm, 1 µm film thickness, J&W Scientific, Folsom, CA, USA), and helium gas was used at a flow rate of 1.2 mL·min-1. The temperature program started at 35°C for 2 min, then increased at 5°C·min-1 to 250°C, and was held at 250°C for 5 min. The mass spectra were obtained with ionization energy of 70 eV, a scan rate of 7 scans·s-1, and a mass-to-charge ratio scan range of 35 to 220. Volatile compounds were tentatively identified through a comparison (concordance > 70%) of the experimental MS spectra and spectra from the mass spectral library of the Wiley Registry (11th edition/NIST 2017). Chromatograms were analyzed using MSD ChemStation data analysis software (Agilent, Santa Clara, CA, USA).

Data analysis

The experimental plots were laid out into five blocks based on a randomized complete block design for each strawberry cultivar. Three strawberry fruits at each stage were randomly collected from the cultivar and analyzed (n = 3). The mean and standard deviation of all data were calculated using the MS Excel program (Microsoft, Redmond, WA, USA). All figures were created using Sigmaplot 14.0 (Systat Software, Inc.).

Results and Discussion

Analysis of volatile compounds by cultivar and maturity stage

The volatile compounds of the four strawberry cultivars (‘Sulhyang’, ‘Maehyang’, ‘Keumsil’, and ‘Arihyang’) were detected by means of GC–MS at different maturation stages (Fig. 2). Consequently, 12 to 59 volatile compounds were qualitatively detected in all cultivars and at all stages (Suppl. Fig. S1-S4). More than 90 volatile compounds have been identified in strawberries, of which 44 contribute to the taste and aroma of strawberries (Du et al., 2011). Strawberry volatile compounds are divided into three categories: sweetness, fruity, and liking (Du et al., 2011; Schwieterman et al., 2014; Fan et al., 2021). The strongest flavor category is fruity, followed by sweetness (Du et al., 2011; Schwieterman et al., 2014; Fan et al., 2021). Fifty-nine volatile compounds were analyzed in four cultivars, among which methyl hexanoate, hexyl acetate, and 4-methoxy-2,5-dimethyl-3(2H)-furanone are related to sweetness, while 1-hexanol and ethyl hexanoate were reported to be related to liking (Du et al., 2011; Schwieterman et al., 2014; Fan et al., 2021).

https://static.apub.kr/journalsite/sites/kshs/2024-042-03/N0130420309/images/HST_42_03_09_F2.jpg
Fig. 2.

GC-MS chromatogram of volatile compounds in the red stage sample extract of 'Sulhyang'. 1: methyl butanoate, 2: methyl isovalerate, 3: hexanal, 4: 2-hexenal, 5: 2-hexen,-1-ol, (e)-, 6: 1-hexanol, 7: methyl hexanoate, 8: ethyl hexanoate, 9: hexyl acetate, 10: 2-ethyl-1-hexanol, 11: 4-methoxy-2,5-dimethyl-3(2h)-furanone, 12: linalool, 13: γ-dodecalactone.

In the ‘Sulhyang’ cultivar, 41, 47, 31, and 53 volatile compounds were detected in the green, white, pink, and red stages, respectively, while methyl isovalerate was detected only in ‘Sulhyang’ (Table 2 and Suppl. Fig. S1). Among these, four, five, seven, and 13 volatile compounds contributing to the taste and aroma of strawberries were identified in the green, white, pink, and red stages, respectively (Table 2 and Suppl. Figs. S1-S4). Methyl isovalerate, closely related to apple and pineapple flavor (Du et al., 2011) and which has been reported to contribute to fruit quality significantly (Campbell et al., 2020), was detected in ‘Sulhyang’.

In the ‘Maehyang’ cultivar, 15, 12, 42, and 55 volatile compounds were detected in the green, white, pink, and red stages, respectively, with the highest relative level of hexyl acetate being found in ‘Maehyang’ (Table 2 and Suppl. Fig. S2). In the ‘Keumsil’ cultivar, 22, 23, 29, and 55 volatile compounds were detected in the green, white, pink, and red stages, respectively (Table 2 and Suppl. Fig. S3). In the ‘Arihyang’ cultivar, 15, 20, 35, and 59 volatile compounds were detected in the green, white, pink, and red stages, respectively, and hexyl butanoate was detected only in ‘Arihyang’ (Table 2 and Suppl. Fig. S4). Hexyl acetate, reportedly associated with fruitiness and sweetness (Schwieterman et al., 2014; Fan et al., 2021), was detected in ‘Maehyang’ (Table 2), whereas hexyl butanoate, which has been reported to be associated with sweetness and overall liking (Schwieterman et al., 2014), was detected in ‘Arihyang’ (Table 2).

In all cultivars, more volatile compounds were detected in the pink and red stages than in the green and white stages (Table 3 and Supplementary row data). Additionally, ten volatile compounds (methyl butanoate; methyl isovalerate; ethyl butyrate; butyl acetate; methyl hexanoate; methyl hexanoate; ethyl hexanoate; 4-methoxy-2,5-dimethyl-3(2H)-furanone; hexyl butyrate; γ-dodecalactone) were detected only in the red stage (Table 2). At the green stage, 15–41 volatile compounds were detected for each cultivar, with ‘Sulhyang’ highest at 41. Among them, there were a total of five major volatile compounds (hexanal; 2-hexenal; 2-Hexen,-1-ol, (E)-; 1-hexanol; 1-hexanol, 2-ethyl-), with ‘Keumsil’ containing the most at five and ‘Arihyang’ containing the least at two (Suppl. Fig. S1-S4). The green stage showed a relatively high level of 1-hexanol compared to the other stages. In the white stage, 12 to 47 volatile compounds were detected for each cultivar, and like the green stage, ‘Sulhyang’ contained the most at 47. There were six major volatile compounds (hexanal; 2-hexenal; 2-hexen,-1-ol, (E)-; 1-hexanol; 1-hexanol, 2-ethyl-), and ‘Arihyang’ was detected least, in only two (Suppl. Fig. S1-S4). In the white stage, 2-hexen,-1-ol, (E)- was increased in the remaining cultivars, except in ‘Sulhyang’; it then tended to decrease in the pink and red stages. At the pink stage, 29 to 42 volatile compounds were detected for each cultivar, with ‘Maehyang’ having the most with 42 detected. There were nine major volatile compounds (methyl butanoate; hexanal; 2-hexenal; 2-hexen,-1-ol, (E)-; 1-hexanol; ethyl hexanoate; hexyl acetate; 1-hexanol,2-ethyl-; linalool), and all nine were detected in ‘Arihyang’ (Suppl. Fig. S1-S4). Specifically, the relative level of hexyl acetate was highest in the pink stage of all cultivars but tended to decrease in the red stage. In the red stage, 53-59 volatile compounds were detected for each cultivar, with ‘Arihyang’ being detected most at 59. There were 17 major volatile compounds (methyl butanoate; methyl isovalerate; hexanal; ethyl butyrate; butyl acetate; 2-hexenal; 2-hexen,-1-ol, (E)-; 1-hexanol; methyl hexanoate; ethyl hexanoate; hexyl acetate; 1-hexanol, 2-ethyl-; 4-methoxy-2,5-dimethyl-3(2H)-furanone; linalool; hexyl butyrate; 1,6,10-dodecatrien-3-ol, 3,7,11-trimethyl-, (E)-; γ-dodecalactone), and 15 were detected in ‘Arihyang’ (Table 2 and Suppl. Fig. S1-S4). When comparing all stages, the red stage exhibited the highest number of volatile compounds. Additionally, methyl butanoate was specifically detected only in ‘Sulhyang’, butyl acetate only in’ Maehyang’, and hexyl butyrate only in ‘Arihyang’. It was previously reported that ten volatile compounds (methyl butanoate; methyl isovalerate; ethyl butyrate; butyl acetate; methyl hexanoate; methyl hexanoate; ethyl hexanoate; 4-methoxy-2,5-dimethyl-3(2H)-furanone; hexyl butyrate; γ-dodecalactone) were detected only in the red stage of all strawberry cultivars, and most of them were related to sweetness (Du et al., 2011; Schwieterman et al., 2014; Fan et al., 2021). Looking at ‘Sulhyang’, seven volatile compounds were detected only in the red stage; accordingly, it is believed that these compounds may be involved in sensory preference (Table 3). However, for more accurate results, a quantitative volatile compound analysis and a sensory evaluation will be necessary.

Table 2.

Odor thresholds and relative levels of volatile compounds contributing to strawberry taste and aroma analyzed in the red stage of four strawberry cultivars

Retention
time
Compound Odor threshold
values
(µg/kg)z
Odor
descriptiony
Relative level
‘Sulhyang’ ‘Maehyang’ ‘Keumsil’ ‘Arihyang’
3.326 Methyl
butanoate
0.01 Sour,
cheesy
28.20 ± 2.83y 5.21 ± 1.72 0.18w NDv NAu 21.31 ± 5.99 0.76
4.406 Methyl
isovalerate
- Fruity,
apple,
pineapple
6.01 ± 0.13 ND NAv ND NA ND NA
4.936 Hexanal - Fresh,
fruity,
green
15.18 ± 2.18 22.00 ± 8.75 1.45 8.92 ± 3.32 0.59 16.15 ± 6.50 1.06
5.018 Ethyl
butanoate
0.00001 Pineapple ND ND NA 15.17 ± 3.21 NA 13.11 ± 6.72 NA
5.425 Butyl
acetate
0.1 Fruity ND 2.47 ± 0.32 NA ND NA ND NA
6.661 2-Hexenal 0.1 Green,
grassy
73.05 ± 11.08 36.95 ± 11.08 0.51 46.64 ± 5.23 0.64 52.08 ± 11.47 0.70
7.197 2-Hexen,
-1-ol, (E)-
0.1 Fruity 82.88 ± 13.18 92.46 ± 18.31 1.12 75.75 ± 5.03 0.91 64.54 ± 3.15 0.78
7.304 1-Hexanol 0.1 Green,
grassy
91.86 ± 14.42 143.68 ± 33.32 1.56 51.63 ± 8.39 0.56 60.51 ± 13.54 0.66
9.809 Methyl
hexanoate
0.1 Fruity,
pineapple,
sweaty,
cheesy
43.22 ± 6.47 3.48 ± 1.12 0.08 24.66 ± 3.76 0.57 40.41 ± 18.54 0.93
13.381 Ethyl
hexanoate
0.0001 Fruity 39.63 ± 19.15 3.59 ± 1.12 0.09 49.00 ± 25.68 1.24 20.01 ± 11.85 0.50
13.916 Hexyl
acetate
0.1 Fruity,
green,
apple,
banana
188.68 ± 39.31 242.45 ± 37.23 1.28 100.67 ± 36.65 0.53 143.41 ± 48.67 0.76
14.455 2-ethyl-
1-Hexanol
- Floral 6.78 ± 0.11 5.94 ± 0.93 0.88 6.17 ± 0.80 0.91 25.60 ± 28.20 3.78
15.479 4-Methoxy-
2,5-dimethyl-
3(2H)-furanone
1.0 Sweet,
caramel
9.68 ± 1.00 2.86 ± 1.30 0.30 7.15 ± 4.03 0.74 22.27 ± 7.39 2.30
16.678 Linalool 0.001 Citrus,
fruity,
floral
6.01 ± 1.84 30.30 ± 3.44 5.04 28.38 ± 5.56 4.72 26.30 ± 2.99 4.38
19.123 Hexyl
butanoate
- Fruity,
apricot
ND ND NA ND NA 4.55 ± 1.60 NA
26.441 1,6,10-
Dodecatrien-3-ol,
3,7,11-trimethyl-,
(E)-
- Woody ND ND NA 29.62 ± 10.09 NA 20.32 ± 24.24 NA
28.338 γ-Dodecalactone 0.01 Peach,
sweet
11.68 ± 2.69 ND NA 9.43 ± 0.44 0.81 13.40 ± 3.62 1.15

x Standard deviation.

w Relative ratio with the relative level of 'Sulhyang'.

v Not detected.

u Not applicable.

Table 3.

Odor thresholds and relative levels of volatile compounds contributing to strawberry taste and aroma analyzed in different maturity stages of ‘Sulhyang’

Retention
time
Compound Odor threshold
values
(µg/kg)z
Odor
descriptiony
Relative level
Green White Pink Red
3.326 Methyl
butanoate
0.01 Sour,
cheesy
NDx ND ND 28.20 ± 2.83w
4.406 Methyl
isovalerate
- Fruity,
apple,
pineapple
ND ND ND 6.01 ± 0.13
4.936 Hexanal - Fresh,
fruity,
green
7.50 ± 1.28 8.69 ± 1.77 5.84 ± 2.03 15.18 ± 2.18
5.018 Ethyl
butanoate
0.00001 Pineapple 25.84 ± 5.41 24.70 ± 3.76 33.99 ± 8.19 73.05 ± 11.08
5.425 Butyl
acetate
0.1 Fruity ND ND 81.66 ± 12.83 82.88 ± 13.18
6.661 2-Hexenal 0.1 Green,
grassy
234.21 ± 5.65 293.90 ± 21.32 132.45 ± 16.67 91.86 ± 14.42
7.197 2-Hexen,
-1-ol, (E)-
0.1 Fruity ND ND ND 43.22 ± 6.47
7.304 1-Hexanol 0.1 Green,
grassy
ND ND 4.31 ± 4.20 39.63 ± 19.15
9.809 Methyl
hexanoate
0.1 Fruity,
pineapple,
sweaty,
cheesy
ND 5.09 ± 1.48 284.41 ± 47.12 188.68 ± 39.31
13.381 Ethyl
hexanoate
0.0001 Fruity 6.64 ± 0.80 15.64 ± 0.93 11.03 ± 0.44 6.78 ± 0.11
13.916 Hexyl
acetate
0.1 Fruity,
green,
apple,
banana
ND ND ND 9.68 ± 1.00
14.455 2-ethyl-
1-hexanol
- Floral ND ND ND 6.01 ± 1.84
15.479 4-Methoxy-2,
5-dimethyl-
3(2H)-furanone
1.0 Sweet,
caramel
ND ND ND 28.20 ± 2.83
16.678 Linalool 0.001 Citrus,
fruity,
floral
ND ND ND 6.01 ± 0.13
19.123 Hexyl
butanoate
- Fruity,
apricot
7.50 ± 1.28 8.69 ± 1.77 5.84 ± 2.03 15.18 ± 2.18
26.441 1,6,10-
Dodecatrien-3-ol,
3,7,11-trimethyl-,
(E)-
- Woody 25.84 ± 5.41 24.70 ± 3.76 33.99 ± 8.19 73.05 ± 11.08
28.338 γ-Dodecalactone 0.01 Peach,
sweet
ND ND 81.66 ± 12.83 82.88 ± 13.18

As reported by Fan et al., (2021), 2-hexenol was tested in 158 cultivars at the pink and red stages and found to be negatively correlated with sweetness and liking. In this study, 1-hexanol and 2-ethyl-1-hexanol, which show fruity and liking flavors (Schwieterman et al., 2014), showed higher relative levels in the green, white, and pink than in the red stage (Suppl. Fig. S1-S4). These compounds were identified in ‘Sulhyang’, ‘Maehyang’, and ‘Keumsil’ (Table 2). Because the relative levels of 1-hexanol and 2-ethyl-1-hexanol were higher in the immature residual stage than in the mature red stage, they are expected to show a negative correlation with sweetness (Schwieterman et al., 2014). The specific volatile compounds that contribute to the organoleptic taste and aroma of strawberries have not yet been fully characterized, and their compositions vary according to the cultivar and growth stage. Therefore, additional research on the functional aspects of aromatic compounds detected in strawberries of different cultivars and stages is required.

Larsen et al. (1992) reported odor threshold values (OTVs) of 24 strawberry volatile compounds categorized as esters, alcohols, carbonyls, and acids (Table 2). OTV refers to the minimum concentration of volatile compounds that the human nose can detect, and the lower the OTV, the better the human nose can detect it. Ethyl butanoate had the lowest OTV of 0.00001 (Larsen et al., 1992) and was detected in the red stage of the ‘Keumsil’ and ‘Arihyang’ cultivars (Table 2). Ethyl butanoate exhibits flavors of pineapple, passionfruit, and strawberry and is used to flavor various beverages (Rodriguez-Nogales et al., 2005; Grosso et al., 2013). Ethyl hexanoate has the second lowest OTV of 0.0001 and is the main aromatic substance of pineapple (Wei et al., 2011). Linalool had the third lowest OTV of 0.001, and similar to ethyl hexanoate, linalool was detected in the red stage of all cultivars (Table 2). Ethyl butanoate and ethyl hexanoate are positively correlated with fruity, sweet, and liking flavors (Fan et al., 2021), whereas linalool is associated with citrus, fruity, and floral flavors (Du et al., 2011). These findings suggest that the three low-OTV compounds discussed above are the major volatile compounds of strawberries and have a significant impact on their flavor.

Nine volatile compounds contributing to strawberry taste and aroma were detected in the red stage of all cultivars in this study (Table 2). In addition, three, two, three, and five unique volatile compounds were detected in ‘Sulhyang’, ‘Maehyang’, ‘Keumsil’, and ‘Arihyang’, respectively. Using ‘Sulhyang’ as a reference cultivar, the relative levels of ethyl butanoate, ethyl hexanoate, and linalool were compared (Table 2). Ethyl butanoate was only detected in the ‘Keumsil’ and ‘Arihyang’ cultivars, and the relative level of ethyl hexanoate was highest in ‘Keumsil’ and lowest in ‘Maehyang’ (Table 2). Linalool was four times more abundant than in ‘Sulhyang’ in ‘Keumsil’, ‘Maehyang’, and ‘Arihyang’ cultivars. These results suggest that the ‘Keumsil’ and ‘Arihyang’ cultivars, in which volatile compounds with low OTVs were detected, can be expected to have a stronger aroma compared to other cultivars. A quantitative analysis of the volatile compounds in strawberries would enable more accurate profiling of their flavor and aroma characteristics.

Analysis of volatile compounds in F. vesca

F. vesca cultivars are derived from wild species in Europe, West Asia, and North America (Dyduch-Siemińska et al., 2015). F. vesca has been widely cultivated in European gardens and has high genetic diversity due to its extensive cultivation range and long cultivation history (Dyduch-Siemińska et al., 2015). F. vesca was evaluated using the ‘Yellow Wonder’ (YW) cultivar with yellow fruits and ‘Baron Solemacher’ (BS) with red fruits, with 53 and 44 volatile compounds detected in YW and BS, respectively (Table 4). Among these, eleven volatile compounds (methyl butanoate; hexanal; ethyl butanoate; butyl acetate; 2-hexanal; 1-hexanol; methyl hexanoate; ethyl hexanoate; hexyl acetate; 2-ethyl-1-hexanol, hexyl butanoate) were associated with strawberry sensory taste and flavor (Du et al., 2011; Schwieterman et al., 2014; Fan et al., 2021). Eleven from VY and eight from BS were detected (Table 3). However, other studies found more than 100 volatile compounds in F. vesca (Ulrich et al., 2007; Urrutia et al., 2017). Differences in the relative levels of aromatic compounds identified in this study can be attributed to environmental factors that arise during cultivation. The relative level of ethyl butanoate was found to be approximately 1.14 times higher in BS than in YW. However, the relative level of ethyl hexanoate was found to be approximately 1.13 times higher in YW than in BS. Compared with domestic strawberry cultivars, ethyl butanoate from the BS cultivar had a relative level that was 8.29–9.60 times higher and ethyl hexanoate had a relative level that was 0.93–10.28 times higher (Fig. 3). Specifically, five volatile compounds (2-hexen,-1-ol, (E)-; 4-methoxy-2,5-dimethyl-3(2H)-furanone; linalool; 1, 6,10-dodecatrien-3-ol, 3,7,11-trimethyl-,(E)-; γ-dodecalactone) detected in four domestic strawberry cultivars were not detected in F. vesca. These results suggest that wild strawberry cultivars are more aromatic than domestic strawberry cultivars.

Table 4.

Odor thresholds and relative levels of volatile compounds contributing to strawberry taste and aroma analyzed in two wild strawberry cultivars

Retention
time
Compound Odor threshold
values (µg/kg)z
Odor descriptiony Relative level
‘Yellow Wonder’ ‘Baron Solemacher’
3.331 Methyl butanoate 0.01 Sour, cheesy 16.17 ± 2.80x 7.35 ± 2.18
4.941 Hexanal - Fresh, fruity, green 12.40 ± 4.56 NDz
5.018 Ethyl butanoate 0.00001 Pineapple 110.58 ± 45.10 125.80 ± 59.00
5.420 Butyl acetate 0.1 Fruity 19.56 ± 1.57 14.27 ± 3.54
6.619 2-Hexanal 0.1 Green, grassy 38.20 ± 13.32 27.15 ± 26.77
7.313 1-Hexanol 0.1 Green, grassy 113.58 ± 34.41 70.39 ± 54.48
9.840 Methyl hexanoate 0.1 Fruity, pineapple,
sweaty, cheesy
10.97 ± 5.47 12.39 ± 8.44
13.398 Ethyl hexanoate 0.0001 Fruity 41.86 ± 30.77 118.86 ± 109.21
13.925 Hexyl acetate 0.1 Fruity, green, apple, banana 155.23 ± 17.62 247.33 ± 160.63
14.468 2-Ethyl-1-hexanol - Floral 3.97 ± 0.61 ND
19.119 Hexyl butanoate - Fruity, apricot 14.82 ± 4.77 ND

https://static.apub.kr/journalsite/sites/kshs/2024-042-03/N0130420309/images/HST_42_03_09_F3.jpg
Fig. 3.

Comparison of relative levels of volatile compounds contributing to strawberry taste and aroma in six strawberry cultivar mature stages: (A) ‘Sulhyang’, (B) ‘Maehyang’, (C) ‘Keumsil’, (D) ‘Arihyang’, (E) ‘Yellow Wonder’, and (F) ‘Baron Solemacher’. 1: methyl butanoate, 2: methyl isovalerate, 3: hexanal, 4: ethyl butyrate, 5: butyl acetate, 6: 2-hexenal, 7: 2-hexen,-1-ol, (e)-, 8: 1-hexanol, 9: methyl hexanoate, 10: ethyl hexanoate, 11: hexyl acetate, 12: 2-ethyl-1-hexanol, 12: 4-methoxy-2,5-dimethyl-3(2h)-furanone, 14: linalool, 15: hexyl butyrate, 16: 1,6,10-dodecatrien-3-ol, 3,7,11-trimethyl-,(e)-, 17: γ-dodecalactone.

Conclusion

Most strawberry cultivars have been developed to produce large, firm fruits with high yields, but fruit aroma is one of the most important characteristics for consumers. If we detect aromatic compounds that exist or that are highly expressed only in cultivars that are strongly preferred by consumers and apply those compounds to breeding, it will then be possible to breed varieties that reflect consumer preferences. Moreover, if the breeding direction is determined by selecting target volatile compounds (methyl butanoate; methyl isovalerate; ethyl butyrate; butyl acetate; methyl hexanoate; methyl hexanoate; ethyl hexanoate; 4-methoxy-2,5-dimethyl-3(2H)-furanone; hexyl butyrate; γ-dodecalactone) based on OTV, breeding will be more efficient. Here, we suggest that ten volatile compounds reported to be related to sweet flavor at the red stage can be used for breeding. However, due to the diversity of strawberry cultivation facilities in Korea, future research should concentrate on the composition of aromatic compounds according to the cultivation environment. Additionally, research on volatile compounds in global domestic strawberry varieties is lacking. Therefore, the qualitative analysis results of the volatile compounds detected in this study can serve as a basis for the future breeding of strawberry fruits with different flavors.

Supplementary Material

Supplementary materials are available at Horticultural Science and Technology website (https://www.hst-j.org).

  • Supplementary Fig. S1. Comparison of peak areas of volatile compounds contributing to strawberry taste and aroma according to fruit development and ripening in ‘Sulhyang’. Each vertical bar represents mean ± SD of three replicates. 1: Methyl butanoate, 2: Methyl isovalerate, 3: Hexanal, 4: Ethyl butyrate, 5: Butyl acetate, 6: 2-Hexenal, 7: 2-Hexen,-1-ol, (E)-, 8: 1-Hexanol, 9: Methyl hexanoate, 10: Ethyl hexanoate, 11: Hexyl acetate, 12: 2-Ethyl-1-Hexanol, 12: 4-Methoxy-2,5-dimethyl-3(2H)-furanone, 14: Linalool, 15: Hexyl butyrate, 16: 1,6,10-Dodecatrien-3-ol, 3,7,11-trimethyl-, (E)-, 17: γ-Dodecalactone.

    HORT_20240028_Fig_S1.pdf
  • Supplementary Fig. S2. Comparison of peak areas of volatile compounds contributing to strawberry taste and aroma according to fruit development and ripening in ‘Maehyang’. Each vertical bar represents mean ± SD of three replicates. 1: Methyl butanoate, 2: Methyl isovalerate, 3: Hexanal, 4: Ethyl butyrate, 5: Butyl acetate, 6: 2-Hexenal, 7: 2-Hexen,-1-ol, (E)-, 8: 1-Hexanol, 9: Methyl hexanoate, 10: Ethyl hexanoate, 11: Hexyl acetate, 12: 2-Ethyl-1-Hexanol, 12: 4-Methoxy-2,5-dimethyl-3(2H)-furanone, 14: Linalool, 15: Hexyl butyrate, 16: 1,6,10-Dodecatrien-3-ol, 3,7,11-trimethyl-, (E)-, 17: γ-Dodecalactone.

    HORT_20240028_Fig_S2.pdf
  • Supplementary Fig. S3. Comparison of peak areas of volatile compounds contributing to strawberry taste and aroma according to fruit development and ripening in ‘Keumsil’. Each vertical bar represents mean ± SD of three replicates. 1: Methyl butanoate, 2: Methyl isovalerate, 3: Hexanal, 4: Ethyl butyrate, 5: Butyl acetate, 6: 2-Hexenal, 7: 2-Hexen,-1-ol, (E)-, 8: 1-Hexanol, 9: Methyl hexanoate, 10: Ethyl hexanoate, 11: Hexyl acetate, 12: 2-Ethyl-1-Hexanol, 12: 4-Methoxy-2,5-dimethyl-3(2H)-furanone, 14: Linalool, 15: Hexyl butyrate, 16: 1,6,10-Dodecatrien-3-ol, 3,7,11-trimethyl-, (E)-, 17: γ-Dodecalactone.

    HORT_20240028_Fig_S3.pdf
  • Supplementary Fig. S4. Comparison of peak areas of volatile compounds contributing to strawberry taste and aroma according to fruit development and ripening in ‘Arihyang’. Each vertical bar represents mean ± SD of three replicates. 1: Methyl butanoate, 2: Methyl isovalerate, 3: Hexanal, 4: Ethyl butyrate, 5: Butyl acetate, 6: 2-Hexenal, 7: 2-Hexen,-1-ol, (E)-, 8: 1-Hexanol, 9: Methyl hexanoate, 10: Ethyl hexanoate, 11: Hexyl acetate, 12: 2-Ethyl-1-Hexanol, 12: 4-Methoxy-2,5-dimethyl-3(2H)-furanone, 14: Linalool, 15: Hexyl butyrate, 16: 1,6,10-Dodecatrien-3-ol, 3,7,11-trimethyl-, (E)-, 17: γ-Dodecalactone.

    HORT_20240028_Fig_S4.pdf

Acknowledgements

This research was funded by the National Research Foundation of Korea (Grant number 2019R1I1A3A01063693).

Disclosure Statement

The authors declare that they have no conflicts of interest.

Author Contributions

Conceptualization, J.M.L. and S.K.K.; writing-original draft preparation, J.S.J. methodology and investigation, H.J.Y. and H.S.S.; data analysis, J.S.J. review and editing, supervision, J.M.L. and S.K.K; funding acquisition, S.K.K. All authors read and approved the manuscript before submitting it.

References

1

Alvarez-Suarez JM, Mazzoni L, Forbes-Hernandez TY, Gasparrini M, Sabbadini S, Giampieri F (2014) The effects of pre-harvest and post-harvest factors on the nutritional quality of strawberry fruits: A review. J Berry Res 4:1-10. doi:10.3233/JBR-140068

10.3233/JBR-140068
2

Arroyo FT, Moreno J, Daza P, Boianova L, Romero F (2007) Antifungal activity of strawberry fruit volatile compounds against Colletotrichum acutatum. J Agric Food Chem 55:5701-5707. doi:10.1021/jf0703957

10.1021/jf070395717567029
3

Campbell SM, Sims CA, Bartoshuk LM, Colquhoun TA, Schwieterman ML, Folta KM (2020) Manipulation of sensory characteristics and volatile compounds in strawberry fruit through the use of isolated wavelengths of light. J Food Sci 85:771-780. doi:10.1111/1750-3841.15044

10.1111/1750-3841.1504432043600
4

Christensen CM (1983) Effects of color on aroma, flavor and texture judgements of foods. J Food Sci 48:787-790. doi:10.1111/j.1365-2621.1983.tb14899.x

10.1111/j.1365-2621.1983.tb14899.x
5

Du X, Plotto A, Baldwin E, Rouseff R (2011) Evaluation of volatiles from two subtropical strawberry cultivars using GC-olfactometry, GC-MS odor activity values, and sensory analysis. J Agric Food Chem 59:12569-12577. doi:10.1021/jf2030924

10.1021/jf203092422026593
6

Dyduch-Siemińska M, Najda A, Dyduch J, Gantner M, Klimek K (2015) The content of secondary metabolites and antioxidant activity of wild strawberry fruit (Fragaria vesca L.). J Anal Methods Chem 2015:831238 doi:10.1155/2015/831238

10.1155/2015/83123826539306PMC4619949
7

Fait A, Hanhineva K, Beleggia R, Dai N, Rogachev I, Nikiforova VJ, Ferhie AR, Aharoni A (2008) Reconfiguration of the achene and receptacle metabolic networks during strawberry fruit development. Plant Physiol 148:730-750. doi:10.1104/pp.108.120691

10.1104/pp.108.12069118715960PMC2556830
8

Fan Z, Hasing T, Johnson TS, Garner DM, Schwieterman ML, Barbey CR, Colquhoun TA, Sims CA, Resende MFR, et al. (2021) Strawberry sweetness and consumer preference are enhanced by specific volatile compounds. Hortic Res 8:224. doi:10.1038/s41438-021-00664-2

10.1038/s41438-021-00664-234615848PMC8494890
9

Food and Agriculture Organization of the United Nations (FAOSTAT) (2021) https://www.fao.org/faostat/en/#data Accessed 7 September 2022

10

Forney CF, Kalt W, Jordan MA (2000) The composition of strawberry aroma is influenced by cultivar, maturity, and storage. HortScience 35:1022-1026. doi:10.21273/HORTSCI.35.6.1022

10.21273/HORTSCI.35.6.1022
11

Giovannoni J (2001) Molecular biology of fruit maturation and ripening. Annu Rev Plant Physiol Plant Mol Biol 52:725-749. doi:10.1146/annurev.arplant.52.1.725

10.1146/annurev.arplant.52.1.72511337414
12

Giovannoni JJ (2004) Genetic regulation of fruit development and ripening. Plant Cell 16:S170-S180. doi:10.1105/tpc.019158

10.1105/tpc.01915815010516PMC2643394
13

Grosso C, Ferreira-Dias S, Pires-Cabral P (2013) Modelling and optimization of ethyl butyrate production catalysed by Rhizopus oryzae lipase. J Food Eng 115:475-480. doi:10.1016/j.jfoodeng.2012.08.001

10.1016/j.jfoodeng.2012.08.001
14

Halford NG, Curtis TY, Muttucumaru N, Postles J, Mottram DS (2011) Sugars in crop plants. Ann Appl Biol 158:1-25. doi:10.1111/j.1744-7348.2010.00443.x

10.1111/j.1744-7348.2010.00443.x
15

Hall RL (1968) Flavor and flavoring seeking a concensus of definition. Food Technology 22:1496

16

Hancock JF (2020) Strawberries (2nd Edition). CABI, Wallingford, UK

17

Karki S, Basak JK, Paudel B, Deb NC, Kim NE, Kook J, Kang MY, Kim HT (2024) Classification of strawberry ripeness stages using machine learning algorithms and colour spaces. Hortic Environ Biotechnol 65:337-354. doi:10.1007/s13580-023-00559-2

10.1007/s13580-023-00559-2
18

Kim SK, Bae RN, Na H, Ko KD, Chun C (2013) Changes in physicochemical characteristics during fruit development in June-bearing strawberry cultivars. Hortic Environ Biotechnol 54:44-51. doi:10.1007/s13580-013-0166-z

10.1007/s13580-013-0166-z
19

Kim SK, Kim DS, Kim DY, Chun C (2015) Variation of bioactive compounds content of 14 oriental strawberry cultivars. Food Chem 184:196-202. doi:10.1016/j.foodchem.2015.03.060

10.1016/j.foodchem.2015.03.06025872444
20

Kim TI, Jang WS, Choi JH, Nam MH, Kim WS, Lee SS (2004) Breeding of Strawberry 'Maehyang' for Forcing Culture. Korean J Hortic Sci Technol 22:434-437

21

Kim TI, Jang WS, Nam MH, Lee WK, Lee SS (2006) Breeding of strawberry 'Sulhyang' for forcing culture. IHC 2006:231

22

Korean Statistical Information Service (KOSIS) (2022) https://kosis.kr/index/index.do Accessed 7 September 2022

23

Larsen M, Poll L, Olsen CE (1992) Evaluation of the aroma composition of some strawberry (Fragaria ananassa Duch) cultivars by use of odour threshold values. Z Lebensm Unters Forsch 195:536-539. doi:10.1007/BF01204558

10.1007/BF01204558
24

Lee CG, Jang PH, Seo JB, Shin GH, Yang WM (2014) A new strawberry 'Jukhyang' with high sugar content and firmness. Acta Hortic 1117:39-44. doi:10.17660/ActaHortic.2016.1117.7

10.17660/ActaHortic.2016.1117.7
25

Lee JM, Kim SK, Lee GD (2003) Monitoring on alcohol fermentation characteristics of strawberry. J Korean Soc Food Sci Nutr 32:679-683. doi:10.3746/jkfn.2003.32.5.679

10.3746/jkfn.2003.32.5.679
26

Li D, Zhang X, Li L, Aghdam MS, Wei X, Liu J, Xu Y, Luo Z (2019) Elevated CO2 delayed the chlorophyll degradation and anthocyanin accumulation in postharvest strawberry fruit. Food Chem 285:163-170. doi:10.1016/j.foodchem.2019.01.150

10.1016/j.foodchem.2019.01.15030797331
27

Menager I, Jost M, Aubert C (2004) Changes in physicochemical characteristics and volatile constituents of strawberry (Cv. Cigaline) during maturation. J Agric Food Chem 52:1248-1254. doi:10.1021/jf0350919

10.1021/jf035091914995129
28

Moya-León MA, Mattus-Araya E, Herrera R (2019) Molecular events occurring during softening of strawberry fruit. Front Plant Sci 10:615. doi:10.3389/fpls.2019.00615

10.3389/fpls.2019.0061531156678PMC6529986
29

Prescott JJ (2004) Psychological processes in flavour perception. Flavor Perception, pp 256-277. doi:10.1002/9780470995716.ch9

10.1002/9780470995716.ch9
30

Rambla JL, Medina A, Fernández-del-Carmen A, Barrantes W, Grandillo S, Cammareri M, Lopez-Casado G, Rodrigo G, Alonso A, et al. (2017) Identification, introgression, and validation of fruit volatile QTLs from a red-fruited wild tomato species. J Exp Bot 68:429-442. doi:10.1093/jxb/erw455

10.1093/jxb/erw45528040800PMC5444475
31

Rodriguez-Nogales JM, Roura E, Contreras E (2005) Biosynthesis of ethyl butyrate using immobilized lipase: a statistical approach. Process Biochem 40:63-68. doi:10.1016/j.procbio.2003.11.049

10.1016/j.procbio.2003.11.049
32

Rural Development Administration (RDA) (2022) https://www.nihhs.go.kr/usr/nihhs/news_Press_view.do Accessed 04 January 2022

33

Schieberle P, Hofmann T (1997) Evaluation of the character impact odorants in fresh strawberry juice by quantitative measurements and sensory studies on model mixtures. J Agric Food Chem 45:227-232. doi:10.1021/jf960366o

10.1021/jf960366o
34

Schwieterman ML, Colquhoun TA, Jaworski RA, Bartoshuk LM, Gilbert JL, Tieman DM, Odabasi AZ, Moskowitz HR, Folta KM, et al. (2014) Strawberry flavor: diverse chemical compositions, a seasonal influence, and effects on sensory perception. PLoS ONE 9:e88446. doi:10.1371/journal.pone.0088446

10.1371/journal.pone.008844624523895PMC3921181
35

Ulrich D, Hoberg E, Rapp A, Kecke S (1997) Analysis of strawberry flavour -discrimination of aroma types by quantification of volatile compounds. Z Lebensm Unters Forsch A Food Res Technol 205:218-223. doi:10.1007/s002170050154

10.1007/s002170050154
36

Ulrich D, Kecke S, Olbricht K (2018) What do we know about the chemistry of strawberry aroma? J Agric Food Chem 66:3291-3301. doi:10.1021/acs.jafc.8b01115

10.1021/acs.jafc.8b0111529533612
37

Ulrich D, Komes D, Olbricht K, Hoberg E (2007) Diversity of aroma patterns in wild and cultivated Fragaria accessions. Genet Resour Crop Evol 54:1185-1196. doi:10.1007/s10722-006-9009-4

10.1007/s10722-006-9009-4
38

Urrutia M, Rambla JL, Alexiou KG, Granell A, Monfort A (2017) Genetic analysis of the wild strawberry (Fragaria vesca) volatile composition. Plant Physiol Biochem 121:99-117. doi:10.1016/j.plaphy.2017.10.015

10.1016/j.plaphy.2017.10.01529100102
39

Vallarino JG, de Abreu e Lima F, Soria C, Tong H, Pott DM, Willmitzer L, Fernie AR, Nikoloski Z, Osorio S (2018) Genetic diversity of strawberry germplasm using metabolomic biomarkers. Sci Rep 8:1-13. doi:10.1038/s41598-018-32212-9

10.1038/s41598-018-32212-930258188PMC6158285
40

Wang J, Yang E, Chaurand P, Raghavan V (2021) Visualizing the distribution of strawberry plant metabolites at different maturity stages by MALDI-TOF imaging mass spectrometry. Food Chem 345:128838. doi:10.1016/j.foodchem.2020.128838

10.1016/j.foodchem.2020.12883833341561
41

Wei CB, Liu SH, Liu YG, Lv LL, Yang WX, Sun GM (2011) Characteristic aroma compounds from different pineapple parts. Molecules 16:5104-5112. doi:10.3390/molecules16065104

10.3390/molecules1606510421694674PMC6264279
42

Xu Y, Charles MT, Luo Z, Mimee B, Tong Z, Véronneau PY, Roussel D, Rolland D (2019) Ultraviolet‐C priming of strawberry leaves against subsequent Mycosphaerella fragariae infection involves the action of reactive oxygen species, plant hormones, and terpenes. Plant Cell Environ 42:815-831. doi:10.1111/pce.13491

10.1111/pce.1349130481398
43

Yoon HS, Jin HJ, Oh JY (2020) 'Kuemsil', a strawberry variety suitable for forcing culture. Korean J Breed Sci 52:184-189. doi:10.9787/KJBS.2020.52.2.184

10.9787/KJBS.2020.52.2.184
44

Zhang J, Wang X, Yu O, Tang J, Gu X, Wan X, Fang C (2011) Metabolic profiling of strawberry (Fragaria× ananassa Duch.) during fruit development and maturation. J Exp Bot 62:1103-1118. doi:10.1093/jxb/erq343

10.1093/jxb/erq34321041374
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