Postharvest Biology and Technology 45 (2007) 349–357

Temperature and relative humidity effects on quality, total ascorbic

acid, phenolics and flavonoid concentrations,
and antioxidant activity of strawberry
Youngjae Shin a,b , Rui Hai Liu a , Jacqueline F. Nock c ,
Darryl Holliday a , Christopher B. Watkins c,∗
aDepartment of Food Science, Cornell University, Ithaca, NY 14853, United States
b Ottogi Food Research Center, Anyang, Kyeonggi 431-070, South Korea
c Department of Horticulture, Cornell University, Ithaca, NY 14853, United States

Received 5 February 2007; accepted 13 March 2007

Abstract
The physical qualities and antioxidant components of ‘Jewel’ strawberry fruit stored in 75, 85 or 95% relative humidity (RH) at 0.5, 10 and
20 ◦ C for 4 days were studied. Overall fruit quality declined more rapidly at 20 ◦ C, especially at 95% RH. Weight loss of fruit was negligible
for 2 days at all temperatures but it increased at 10 ◦ C in the lowest RH and increased rapidly from day 3 at 20 ◦ C especially with lower RH.
Firmness was maintained, or even increased, at 0.5 or 10 ◦ C, while soluble solids concentrations (SSC) decreased at higher storage temperatures.
Red color, assessed using chroma, hue and lightness, and anthocyanin concentrations were relatively unchanged at 0.5 or 10 ◦ C but increased
rapidly at 20 ◦ C as fruit ripened. Firmness, SSC and color were not affected by RH. Total phenolic compounds were slightly higher at 20 ◦ C than at
other temperatures at all RHs. Total ascorbic acid concentrations of the fruit remained similar for the first 2 days of storage, then declined in fruit
stored at 0.5 and 20 ◦ C, but remained unchanged at 10 ◦ C at all RHs. Total flavonoid content of fruit did not change over time at all temperatures.
The total antioxidant activity of fruit was higher at 10 ◦ C than at 0.5 and 20 ◦ C on day 3, and no effect of RH was detected. In conclusion, while
the best temperature for long-term storage is 0.5 ◦ C, quality could be maintained at 10 ◦ C for acceptable periods of time for marketing and may be
associated with better nutritional quality.
© 2007 Elsevier B.V. All rights reserved.

Keywords: Strawberry; Fragaria × ananassa Duch.; Storage; Quality; Ascorbic acid; Phenolics; Flavonoids; Anthocyanin; Antioxidant activity; Fruit

1. Introduction
Much of the research on maintenance of strawberry (Fra-
garia × ananassa Duch.) quality after harvest has centered on
industries that produce berries over an extended production
period and accordingly have high investments in the infras-
tructure required for longer term storage and export of fruit.
In contrast, strawberry fruit grown in regions such as the North-
east USA have a restricted production period, and retail supply
periods are limited. While cooling of fruit in this region is recom-
mended (Pritts and Watkins, 1998), many growers lack adequate
facilities for storage. Moreover, many New York growers have
observed that cooling of fruit to 0 ◦ C can be detrimental for
∗ Corresponding author.
E-mail address: cbw3@cornell.edu (C.B. Watkins).
short term sales, fruit appearance being duller and condensation
of fruit during re-warming resulting in greater decay incidence.
Some growers have obtained good results on quality mainte-
nance by holding fruit at a moderate temperature (10 ◦ C) under
high relative humidity (RH) (pers. comm.).
Assessment of strawberry quality for the market is focused on
visual and internal characteristics, such as size, color, firmness,
acidity, sweetness and aroma (Azodanlou et al., 2003; Mitcham,
2004), but there is increasing interest on the health benefits of the
fruit. Beneficial effects of strawberries include increased plasma
antioxidant capacity in humans (Cao et al., 1998), antioxidant
activity for low-density lipoproteins (Heinonen et al., 1998), and
anticarcinogenic activity against human and mouse cancer cells
(Smith et al., 2004; Wang et al., 2005). Strawberry fruit have
high ascorbic acid concentrations which have protective roles
against reactive oxygen species (Davey et al., 2000) but it has
become increasingly clear that many other components within

0925-5214/$ – see front matter © 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.postharvbio.2007.03.007
350 Y. Shin et al. / Postharvest Biology and Technology 45 (2007) 349–357
the fruit also have antioxidant activity. These include antho-
cyanins, which increase in concentration during ripening and
are important to fruit color (Cheng and Breen, 1991; Given et
al., 1988). As major contributors to the total phenolic contents
in strawberry, anthocyanins also have potent antioxidant prop-
erties (Wang et al., 1997) and reduce oxidative stress-induced
neurotoxicity (Heo and Lee, 2005).
Reactions among antioxidants may be synergistic and there-
fore total antioxidant activity measurements potentially provide
a better estimate of the overall contributions of antioxidant com-
ponents (Liu, 2003; Vinson et al., 2001; Wang et al., 1996).
Strawberries had the highest oxygen radical absorbance capac-
ity (ORAC) of 12 fruit tested (Wang et al., 1996) and were fourth
of 11 fruit tested using the total oxyradical scavenging capacity
(TOSC) and antiproliferation activity by Sun et al. (2002). Total
phenolic concentrations were positively related to total antiox-
idant activity (Meyers et al., 2003; Rekika et al., 2005; Sun et
al., 2002).
The effect of temperature on quality of strawberries, includ-
ing nutritional aspects, has been studied by several research
groups; however, both nutritional and physical quality data are
not always presented, and no study has involved control of RH.
Kalt et al. (1999) found that ascorbic acid and total phenolic
concentrations were stable in strawberry fruit kept at 0, 10,
20 and 30 ◦ C, but anthocyanin concentrations and total antioxi-
dant activity (ORAC) increased with both temperature and time.
No data on physical fruit quality were provided in that study.
Ayala-Zavala et al. (2004) found that strawberry fruit quality
was maintained longer at 0 ◦ C than at 5 or 10 ◦ C over a 13-
day postharvest period, but the strawberry fruit stored at lower
temperatures had lower antioxidant capacity, total phenolics and
anthocyanin concentrations. While ascorbic acid concentrations
decreased in all cultivars during air storage at 6 ◦ C for 6 days,
anthocyanin concentration increases were affected by cultivar
(Cordenunsi et al., 2003). Effects of cultivar on ascorbic acid
and anthocyanins of fruit at 6, 16 and 25 ◦ C for 6 days were also
shown by Cordenunsi et al. (2005). Total antioxidant activity
decreased during storage and was not affected by cultivar.
RH was not controlled in any of the above studies, and there-
fore the extent to which variable water losses may have affected
the results is not known. Nunes et al. (1998) compared ascorbic
acid changes in fruit at 1, 10 and 20 ◦ C in unwrapped and plastic
wrapped containers. Wrapping reduced ascorbic acid loss, and
the authors concluded that water loss had greater effects on these
losses than temperature.
The objective of our study was to investigate the effects of
temperature under controlled RH conditions on physical and
nutritional status of strawberry fruit harvested at the red ripe
stage of maturity for short postharvest storage periods of up to
4 days.
2. Materials and methods
2.1. Fruit harvest, treatment and storage conditions
Strawberry fruit (Fragaria × ananassas Duch., cv. Jewel)
were harvested at the red ripe stage at a local commercial farm
and transported to the Cornell University Postharvest Labora-
tory. Fruit were then sorted to eliminate damaged fruit, and
selected for uniform size and color.
About 45 fruit were placed in each of 108 1.9 L Mason
jars, which were weighed before and after the addition of fruit.
After allowing fruit temperatures to equilibrate to the appro-
priate storage temperatures (0.5, 10, and 20 ◦ C) for 2 h, the
jars were connected to flow boards established with water and
glycerol mixtures that maintained relative humidities of 75, 85
or 95% (Osborne and Bacon, 1961). The RH of each flow
board was verified several times daily with a Traceable digi-
tal hygrometer/thermometer (Control Company, Friendswood,
TX, USA). The flow rate through each jar was 100 mL min−1 .
Three replicate jars were removed from each temperature and
RH treatment daily for up to 4 days and quality of the fruit was
assessed.
2.2. Weight loss, overall quality, and decay assessments
Fruit weights were recorded, and the percentage weight loss
from harvest was calculated. All fruit were evaluated for qual-
ity on a 1–5 scale according to the percentage of surface area
damaged, where 1 = unacceptable (>50% surface showing skin
damage or discoloration), 2 = bad (20–50% surface affected),
3 = acceptable (5–20% surface affected), 4 = good (up to 5%
surface affected), and 5 = excellent. Results were expressed
as an overall quality index. Fruit with visible mold growth
were expressed as percentage of fruit showing decay symp-
toms.
Fruit, free of excessive damage or decay, were used for
subsequent analyses. Ten fruit were used for physical quality
assessments. The remaining fruit were frozen after they were
sliced into liquid nitrogen and kept at −80 ◦ C until used for
extraction and measurement of total phenolic, flavonoid, antho-
cyanin and ascorbic acid concentrations, and total antioxidant
activity.
2.3. Color, firmness, soluble solid concentration (SSC), pH,
and total titratable acidity (TA)
The fruit surface color was measured using a chromame-
ter (CR 100, Minolta, Ramsey, NJ). The measurements are
expressed as chroma (intensity of color), hue angle (actual color,
or redness), and L* value (dark to light, on a scale of 0–100).
Three readings were taken around the equatorial region of each
fruit and the average of the values for each fruit was calcu-
lated.
Firmness was measured by a puncture test on each fruit with
intact skin using a Force Five pressure tester (Model FDV-30,
Wagner Instruments, Greenwich, CT, USA) fitted with a 7.9 mm
diameter flathead probe.
The fruit were then wrapped in cheesecloth, squeezed by
hand, and the expressed juice used for measurements of SSC,
pH, and TA. SSC was determined at room temperature using an
Atago PR-100 refractometer (Atago Co. Ltd., Tokyo, Japan).
pH and TA were determined with a Mettler DL 12 titrator
(Mettler-Toledo Inc., NJ, USA) after diluting each 5 mL aliquot
 Y. Shin et al. / Postharvest Biology and Technology 45 (2007) 349–357
of strawberry juice in 95 mL of distilled water. Titration was
carried out to pH 8.2 using 0.1 mol L−1 NaOH.
2.4. Extraction and measurement of flavonoids, phenolics
and total antioxidant activity
Phytochemical extracts were extracted from frozen straw-
berry using 80% acetone as described previously (Liu et al.,
2002). The frozen strawberry slices were crushed into coarse
pieces, and 10 g of fruit was homogenized with 100 mL of 80%
acetone using a Waring commercial blender (Black & Decker,
MD, USA) for 3 min. The homogenate was filtered through
#1 Whatman paper. The filtrate was recovered and the acetone
was evaporated by a rotary evaporator at 45 ◦ C for 30 min. The
samples were then brought to the desired volume (10 mL) with
deionized water and kept frozen at −80 ◦ C prior to analysis. The
extracts were used for the measurement of flavonoids, phenolics,
and total antioxidant activity.
The total flavonoid contents were determined by a colorimet-
ric assay (Jia et al., 1999; Meyers et al., 2003). A 1 mL aliquot of
appropriately diluted sample was added to a 15 mL tube contain-
ing 4 mL of deionized water. Then 0.3 mL of 5% NaNO2 was
added to this mixture, which was allowed to stand for 5 min at
room temperature, and 0.3 mL of 10% AlCl3 ·6H2 O was added.
The mixture was allowed to stand for 6 min at room temperature,
and 2 mL of 1 mol L−1 NaOH was added, and the total was made
up to 10 mL with deionized water. The absorbance of the solu-
tion versus a blank at 510 nm was measured immediately. The
results are expressed as catechin equivalents using a standard
curve prepared from authentic catechin.
The total phenolic contents of the strawberry extracts were
measured using a modified Folin-Ciocalteu colorimetric method
(Meyers et al., 2003; Singleton et al., 1999). A 0.2 mL sample
of the extract was added to a 15 mL tube containing 2.6 mL of
deionized water, 0.2 mL of Folin-Ciocalteu reagent was added
to the mixture, and the tube was stoppered and allowed to stand
at room temperature for 6 min. Then, 2 mL of 7% Na2 CO3 was
added to the mixture. Absorbance was measured at 750 nm ver-
sus a blank after 90 min at room temperature. The results are
expressed as gallic acid equivalent on a fresh weight basis,
mg kg−1 .
The total antioxidant activity of strawberry extracts of fruit at
harvest and after 3 days of storage was measured using the total
oxyradical scavenging capacity (TOSC) assay (Winston et al.,
1998) as modified in our lab (Dewanto et al., 2002). Antioxidant
activity was assessed at four different time points (15, 30, 45,
and 60 min) and six concentrations to determine the TOSC value.
The TOSC value was quantified from the integration of the area
under the kinetic curve according to the equation:
  3.1. Overall quality
SA
TOSC = 100 −  × 100
CA
 
where SA and CA are the integrated areas from the sam-
ple and the control reaction, respectively. The results were
calculated as vitamin C equivalents on a fresh weight basis,
mmol kg−1 (Dewanto et al., 2002).
351
2.5. Extraction and measurement of total anthocyanin
concentrations
Five grams of frozen tissue was ground to a fine powder
under liquid nitrogen by cold mortar and pestle and 1 g of the
resultant powder was added to 10 mL of methanol containing
HCl (1%, v/v) and held at 0 ◦ C for 10 min (Cordenunsi et al.,
2003). The slurry was centrifuged at 17,600 × g for 15 min at
4 ◦ C, the supernatant was used and its absorbance at 515 nm was
measured. The amount of anthocyanins was calculated using
the extinction coefficient (ε) equal to 3.6 × 106 mol−1 m−1
(Woodward, 1977). Total anthocyanin content was expressed
as pelargonidin-3-glucoside on a fresh weight basis,
mg kg−1 .
2.6. Extraction and measurement of total ascorbate
concentrations
Total ascorbic acid (AA) content was determined using the
dinitrophenylhydrazine (DNPH) method (Terada et al., 1978).
Five grams of homogenized fruit tissue was added to 100 mL
of a mixture of 6% metaphosphoric acid in 2 mol L−1 acetic
acid. The mixture was centrifuged at 17,600 × g for 15 min at
4 ◦ C and the supernatant was filtered through #1 Whatman fil-
ter paper. A 1 mL aliquot of the supernatant was mixed with
0.05 mL of 0.2% 2,6-dichlorophenolindolphenol (DCIP) and the
solution was incubated at room temperature for 1 h. After that,
1 mL of 2% thiourea in 5% metaphosphoric acid and 0.5 mL
of 2% DNPH in 4.5 mol L−1 sulfuric acid were added to the
solution, and then incubated at 60 ◦ C for 3 h. The reaction was
stopped by placing the tubes in an ice bath and slowly adding
2.5 mL of ice cold 90% sulfuric acid. Total AA was measured
by absorbance at 540 nm using a standard curve. The concen-
trations were expressed as ascorbic acid on a fresh weight basis,
mg kg−1 .
2.7. Statistical analysis
Data were subjected to analysis of variance using the gen-
eral linear model procedure to determine the main effects and
interactions (Minitab Release 13.1, State College, PA). Where
appropriate, further data analyses were carried out within each
temperature regimen. Sources of variation were storage duration
and treatment. Percentage decay data were arc sine transformed
before analysis. Mean comparisons were performed using the
LSD at the 0.05 level.
3. Results
Quality ratings of fruit showed a relatively small change over
time at 0.5 ◦ C, declining from an overall rating of 4.8 at harvest to
4.0 on day 4 (Fig. 1). At 10 ◦ C, ratings declined slightly faster,
declining to 4.1 by day 2 and 3.6 by day 4. No effect of RH
was detected at either 0.5 or 10 ◦ C. At 20 ◦ C, quality remained
relatively high on day 2 (average rating of 4.1), but then declined
352 Y. Shin et al. / Postharvest Biology and Technology 45 (2007) 349–357

Table 1
Fungal decay of ‘Jewel’ strawberry fruit stored for up to 4 days at 0.5, 10 and
20 ◦ C in 75, 85 or 95% RH
Temperature (◦ C) RH (%) Fungal decay (%)

Day 3 Day 4

75 0 (0) 0 (0)
0.5 85 0 (0) 0 (0)
95 0 (0) 0 (0)
75 0 (0) 3.9 (1)
10 85 0 (0) 9.3 (4)
95 0 (0) 11.7 (7)
75 9.3 (4) 59.2 (65)
20 85 20.1 (12) 90.0 (100)
95 39.7 (41) 90.0 (100)
LSD (P = 0.05) 15.77

No decay was detected on either day 1 or 2 of storage. The LSD (P = 0.05)

provided is for comparison of arc sine transformed means. Back-transformed
means are in parentheses.

3.2. Color

Lightness (L* value), intensity of color (C* value) and red-

ness (h◦ value) were not affected by RH and therefore only
changes over time at each temperature are shown (Fig. 3). Light-
ness of the fruit was not affected by storage time at 0.5 ◦ C, but
declined slightly with higher storage temperatures. Chroma and
hue values remained similar at 0.5 ◦ C, but decreased at higher
temperatures (10 and 20 ◦ C) showing the fruit turned redder
(Fig. 3), as would be expected from faster ripening at warm
temperatures.

3.3. Firmness, SSC, TA and pH

Berry firmness was not affected by RH and therefore only

Fig. 1. Quality index of ‘Jewel’ strawberry fruit stored for up to 4 days at 0.5,
10 and 20 ◦ C in 75, 85 or 95% RH. The highest order interaction detected was
for temperature, RH and storage time (P = 0.017). The vertical bar represents
the LSD value at P = 0.05 for comparison of means across all temperatures.
slightly faster to averages of 3.3 on day 3, and then more rapidly
(Fig. 1). Loss of quality was greater at 95% than at 75 or 85%
on day 3, and lowest at 75% on day 4.
Fungal decay was the major contributor to loss of qual-
ity by day 3 at 20 ◦ C, and to a lesser extent at 10 ◦ C, where
decay occurred on day 4 (Table 1). No decay was detected at
0.5 ◦ C.
Weight loss from the berries increased over time (Fig. 2) and
was affected by storage temperature (P < 0.001). Overall weight
losses were similar on days 1 and 2, but by day 4 were 0.42 and
0.61% at 0.5 and 10 ◦ C, respectively; and by day 3 were 0.81%
at 20 ◦ C. Weight loss was unaffected by RH at 0.5 and 10 ◦ C
until day 3 when it was greater at 75% than at 85 and 95% RH.
The effects of RH were more pronounced as storage temperature
increased.
changes over time at each temperature are shown (Fig. 4). At
0.5 and 10 ◦ C, firmness increased from at harvest values on day
2 and beyond. At 20 ◦ C, the fruit first softened but then firm-
ness increased by day 3. The SSC of fruit at harvest was 6.5%
and was affected by storage time (P = 0.002) and temperature
(P = 0.002). At 0.5 ◦ C, the average SSC were 6.7% compared
with 6.5 and 6.4% at 10 and 20 ◦ C, respectively. Overall, con-
centrations declined to 6.3% on day 3. An interaction between
storage time and temperature was detected (P = 0.009), but there
were no consistent patterns of change (data not shown). TA
and pH remained similar, being 0.57% and 3.09 units over all
treatments (data not shown).
3.4. Total ascorbic acid concentration
The total ascorbic acid concentrations of the fruit remained
similar for the first 2 days of storage, irrespective of storage
temperature (Fig. 5). The concentrations were not affected by
RH (P = 0.278). The ascorbic acid concentrations then declined
in fruit stored at 0.5 ◦ C (P = 0.007) and 20 ◦ C (P = 0.002), but
remained unchanged at 10 ◦ C (P = 0.280).
 Y. Shin et al. / Postharvest Biology and Technology 45 (2007) 349–357
Fig. 2. Weight loss (%) of ‘Jewel’ strawberry fruit stored for up to 4 days at
0.5, 10 and 20 ◦ C in 75, 85 or 95 RH. The effects of temperature, storage time,
their interactions (P ≤ 0.001), and RH (P = 0.034) were significant. The vertical
bars represent the LSD values at P = 0.05 for comparison of means within each
temperature.
3.5. Anthocyanin, total flavonoid and total phenolic
contents
The anthocyanin concentrations of the berries were not
affected by RH at any storage temperature (P = 0.105). The
concentrations of anthocyanin increased markedly within a day
at 20 ◦ C (Fig. 6), while those of fruit stored at 0.5 and 10 ◦ C
remained relatively unchanged. Overall, those concentrations
averaged 87.2 and 99.7 mg kg−1 , at 0.5 and 10 ◦ C, respectively,
compared with 136.6 mg kg−1 at 20 ◦ C.
The flavonoid concentrations were affected only by RH
(P = 0.05), averaging 624, 638 and 651 mg kg−1 at 75, 85, and
95% RH, respectively (data not shown).
The total phenolics concentrations were affected by days
(P = 0.001), temperature (P = 0.023) and RH (P = 0.001).
353
Fig. 3. Lightness, chroma and hue values of ‘Jewel’ strawberry fruit stored
for up to 4 days at 0.5, 10 and 20 ◦ C. No effects of RH were detected and
therefore data have been combined for each parameter. For lightness, only effects
of temperature (P ≤ 0.001), and storage time × temperature (P = 0.004) were
significant. For both chroma and hue, effects of temperature, storage time, and
their interactions, were significant (P ≤ 0.001). The vertical bars represent the
LSD value at P = 0.05 for comparison of means within each parameter.
Patterns of change were not consistent (Fig. 7), although the
phenolic concentrations were generally higher at 75 and 85%
RH compared with 95% RH. At harvest, the concentrations
were 2240 mg kg−1 , and over all temperatures and RHs, these
increased to 2480 and 2300 mg kg−1 on days 2 and 3, before
declining. Overall, the highest concentration (2470 mg kg−1 )
occurred at 20 ◦ C, compared with 2310 and 2290 mg kg−1
at 0.5 and 10 ◦ C, respectively. The phenolic concentrations
averaged 2280, 2350 and 2410 mg kg−1 at 75, 85, and 95% RH,
respectively.
354 Y. Shin et al. / Postharvest Biology and Technology 45 (2007) 349–357

Fig. 4. Firmness (N) of ‘Jewel’ strawberry fruit stored for up to 4 days at 0.5,
10 and 20 ◦ C. No effects of RH were detected and therefore data have been
combined for each temperature. The effects of temperature, storage time, and
their interactions, were significant (P ≤ 0.001). The vertical bar represents the
LSD value at P = 0.05 for comparison of means.

Fig. 5. Total ascorbic acid concentrations (mg kg−1 fresh weight) of ‘Jewel’
strawberry fruit stored for up to 4 days at 0.5, 10 and 20 ◦ C. No effects of RH
were detected and therefore data have been combined for each temperature. The
effect of temperature × storage time was significant (P = 0.011). The vertical bar
represents the LSD value at P = 0.05 for comparison of means.

Fig. 7. Total phenolics (gallic acid equivalent on a fresh weight basis, mg kg−1 )
of ‘Jewel’ strawberry fruit stored for up to 4 days at 0.5, 10 and 20 ◦ C in 75, 85
or 95% RH. The highest order interaction detected was for RH and storage time
(P = 0.034). The vertical bar represents the LSD value at P = 0.05 for comparison
of means across all temperatures.

3.6. Total antioxidant activity

The total antioxidant activity was measured only on fruit

Fig. 6. Anthocyanin concentrations (pelargonidin-3-glucoside on a fresh weight
basis, mg kg−1 ) of ‘Jewel’ strawberry fruit stored for up to 4 days at 0.5, 10 and
20 ◦ C. No effects of RH were detected and therefore data have been combined
for each temperature. The effect of temperature was significant (P ≤ 0.001). The
vertical bar represents the LSD value at P = 0.05 for comparison of means.
at harvest and after 3 days of storage. At harvest, the total
antioxidant activity of the berries was 32.5 mmol kg−1 , vita-
min C equivalents on a fresh weight basis. On day 3 of storage
(Table 2), the activity was not affected by RH (P = 0.651),
but was marginally higher (P = 0.052) at 10 ◦ C, averaging
41.9 mmol kg−1 compared with 35.6 and 34.9 mmol kg−1 at 0.5
and 20 ◦ C, respectively.
 Y. Shin et al. / Postharvest Biology and Technology 45 (2007) 349–357
4. Discussion
From a marketing perspective, the critical feature of straw-
berry quality is appearance or visual quality, and our results
agree with the well-known benefit of low storage temperature on
maintaining this aspect of quality (Mitcham, 2004). The results
indicate, however, that different aspects of quality are affected
by these treatment factors to varying degrees during storage.
Storage at 0.5 ◦ C resulted in better visual quality than at
10 C, which was better than 20 ◦ C (Fig. 1). A major cause of
◦
loss of visual quality was the onset of fungal decay (Table 1).
These results are similar to those of Ayala-Zavala et al. (2004)
who found lower storage temperatures positively affected fruit
quality and decreased the fungal decay incidence. Our study,
however, was focused on the effects of storage temperature on
ripe berries that would be marketed soon after harvest, and also
incorporated the effects of RH. The effect of storage temperature
on visual quality was relatively small for the first 2 days after
harvest, particularly at 10 ◦ C, thereby supporting industry obser-
vations that deterioration can be slowed even at a higher than
recommended temperature. No effect of RH on visual quality or
decay was detected at 0.5 ◦ C, but higher RH was associated with
greater loss of quality and increasing decay as storage tempera-
ture increased (Fig. 1 and Table 1). Weight loss was affected by
storage temperature and RH (Fig. 2), but even at the lowest RH
and highest storage temperature on day 3, it was low (1.04%)
compared with that in other studies. Nunes et al. (1998) found
that the weight losses of unwrapped strawberries stored at 20 ◦ C
were about 16 and 23% on days 3 and 4, respectively, and were
four-fold greater than wrapped strawberries.
Red color development remained relatively stable at 0.5 and
10 ◦ C, but at 20 ◦ C, color changed to dark red (Figs. 3 and 6). No
effect of RH was detected. Higher storage temperatures result
in faster ripening and greater anthocyanin accumulation (Ayala-
Zavala et al., 2004; Cordenunsi et al., 2003, 2005; Kalt et al.,
1999; Zhang and Watkins, 2005).
Firmness of the berries increased at 0.5 and 10 ◦ C during
the storage time (Fig. 4). Increasing firmness as a result of
cold-storage is a common phenomenon in fruit and vegetables,
including strawberries (Bourne, 1982; Harker et al., 1997). The
increased firmness might be due to an increase of the pectin
viscosity (the gelling behavior) rather than metabolic processes
(Werner and Frenkel, 1978; Werner et al., 1978). In strawberries,
the firming effect is enhanced by 15–20% CO2 in the storage
atmosphere (Watkins et al., 1999; Zhang and Watkins, 2005),
and is also ascribed to physical rather than metabolic effects
(Harker et al., 2000). The firming effect can vary among culti-
vars (Cordenunsi et al., 2003; Watkins et al., 1999). An increase
in berry firmness, after an initial decline, was also observed in
fruit kept at 20 ◦ C (Fig. 4), even though fruit were visually red-
der (Fig. 3). It is possible that this finding was an artifact; less
berries suitable for analyses were available by day 3 and firmer
fruit may be associated with less decay.
The SSC was reduced at 20 ◦ C compared with the lower stor-
age temperatures, although it was not affected by RH, whereas
TA and pH were not affected by either temperature or RH.
Decreased SSC at higher temperatures is probably due to higher
355
respiration rates (Ayala-Zavala et al., 2004). However, effects of
temperature on SSC can be affected by cultivar (Cordenunsi et
al., 2005). The absence of effects of storage temperature and RH
on TA and pH has also been found by Cordenunsi et al. (2003)
and Ayala-Zavala et al. (2004).
Ascorbic acid has long been considered an important nutri-
tional component of strawberry fruit. Over the short time period
used in our study, the concentrations remained similar at all
storage temperatures for the first 2 days (Fig. 5) and were not
affected by RH. The concentrations then declined at 0.5 and
20 ◦ C. The decline at 20 ◦ C might be associated with loss of
quality (Fig. 1) and faster ripening as indicated by reddening
(Fig. 3), but a decline at 0.5 ◦ C is less easily explained. In gen-
eral, ascorbic acid in fruits is considered to be more stable during
storage compared with leafy products because of their high acid
content. Nevertheless water loss may enhance loss of ascorbic
acid because of increased oxidation (Nunes et al., 1998).
The total flavonoid concentration of strawberry fruit can be
maintained or changed by storage conditions (Cordenunsi et al.,
2005). Our study showed that the flavonoid concentration was
slightly higher at 95% RH than at 75 and 85% RH (data not
shown). The concentration of total phenolics of strawberry fruit
can be maintained or changed during storage (Ayala-Zavala et
al., 2004; Kalt et al., 1999). In our study, the total phenolic
concentration was higher at 20 ◦ C than other storage tempera-
tures over the storage time (Fig. 7). Ayala-Zavala et al. (2004)
found that the total phenolics increased continuously in straw-
berry fruit stored at 10 and 5 ◦ C. However, the total phenolics
of strawberry fruit stored at 0 ◦ C were maintained well over a
13-day postharvest period. The total phenolic contents were sig-
nificantly affected by the storage temperature and storage period.
These results contrast with those of Cordenunsi et al. (2005),
where the total phenolic contents remained or even decreased in
three different cultivars at all temperatures.
The total antioxidant activity was assessed only in samples at
harvest and those from day 3 as fruit quality of samples stored at
20 ◦ C had deteriorated excessively by day 4. In addition, while
the anthocyanin concentrations increased greatly at 20 ◦ C, few
changes in the concentration of total phenolics, total flavonoids
and total ascorbic acids had occurred at the early storage times.
The total antioxidant activity was not affected by the RH but was
highest at 10 ◦ C (41.9 mmol kg−1 ) than at 0.5 and 20 ◦ C (35.6
and 34.9 mmol kg−1 , respectively) on day 3 (Table 2). Kalt et al.
(1999) reported that the total antioxidant capacity of the straw-
berry fruit increased 1.5-fold during the 8 days of storage, with
greater increase occurring at 10 and 20 ◦ C than at 0 ◦ C. Ayala-
Zavala et al. (2004) also found that the total antioxidant capacity
changed very little at 0 ◦ C during storage time but increased
significantly at 5 and 10 ◦ C. Even though the overall quality
was better maintained at 0 ◦ C, the storage temperature of 10 ◦ C
significantly increased the total antioxidant capacity, total antho-
cyanin, and total phenolics (Ayala-Zavala et al., 2004). In the
present study, however, the individual compounds did not have a
relationship with the total antioxidant activity. The total phenolic
and the anthocyanin concentrations were higher at 20 ◦ C than at
0.5 and 10 ◦ C. The total ascorbic acid concentration was higher
at 10 ◦ C, whereas the total flavonoid concentrations were not
356 Y. Shin et al. / Postharvest Biology and Technology 45 (2007) 349–357

Table 2 Azodanlou, R., Darbellay, C., Luisier, J.L., Villettaz, J.C., Amado, R., 2003.
Total oxygen scavenging concentration (vitamin C equivalents on a fresh weight Quality assessment of strawberries (Fragaria species). J. Agric. Food Chem.
basis, mmol kg−1 ) of ‘Jewel’ strawberry fruit stored for 3 days at 0.5, 10 and 51, 715–721.
20 ◦ C in 75, 85 or 95% RH Bourne, M.C., 1982. Effect of temperature on firmness of raw fruits and vegeta-
bles. J. Food Sci. 47, 440–444.
Temperature (◦ C) RH (%) TOSC (mmol kg−1 )
Cao, G.H., Russell, R.M., Lischner, N., Prior, R.L., 1998. Serum antioxidant
Day 3 Temp. capacity is increased by consumption of strawberries, spinach, red wine or
vitamin C in elderly women. J. Nutr. 128, 2383–2390.
75 38.9 Cheng, G.W., Breen, P.J., 1991. Activity of phenylalanine ammonia-lyase (PAL)
0.5 85 32.2 35.6 and concentrations of anthocyanins and phenolics in developing strawberry
95 35.7 fruit. J. Am. Soc. Hort. Sci. 116, 865–869.
75 41.9 Cordenunsi, B.R., Genovese, M.I., Nascimento, J.R.O., Hassimotto, N.M.A.,
10 85 44.3 41.9 Santos, R.J., Lajolo, F.M., 2005. Effects of temperature on the chemical com-
95 39.6 position and antioxidant activity of three strawberry cultivars. Food Chem.
91, 113–121.
75 36.3 Cordenunsi, B.R., Nascimento, J.R.O., Lajolo, F.M., 2003. Physico-chemical
20 85 32.7 34.9 changes related to quality of five strawberry fruit cultivars during cool-
95 36.3 storage. Food Chem. 83, 167–173.
LSD (P = 0.05) 6.33 Davey, M.W., Van Montagu, M., Inze, D., Sanmartin, M., Kanellis, A., Smirnoff,
N., Benzie, I.J.J., Strain, J.J., Favell, D., Fletcher, J., 2000. Plant l-ascorbic
The LSD (P = 0.05) provided is for comparison of means for temperature. acid: chemistry, function, metabolism, bioavailability and effects of process-
ing. J. Sci. Food Agric. 80, 825–860.
Dewanto, V., Wu, X.Z., Adom, K.K., Liu, R.H., 2002. Thermal processing
different for the different storage temperatures on day 3. Several enhances the nutritional value of tomatoes by increasing total antioxidant
studies showed that total phenolic concentrations are positively activity. J. Agric. Food Chem. 50, 3010–3014.
related to total antioxidant activity (Meyers et al., 2003; Rekika Given, N.K., Venis, M.A., Grierson, D., 1988. Phenylalanine ammonia-lyase
et al., 2005; Sun et al., 2002) but we did not find relationships activity and anthocyanin synthesis in ripening strawberry fruit. J. Plant
Physiol. 133, 25–30.
over a smaller range of phenolic concentration than described Harker, F.R., Elgar, H.J., Watkins, C.B., Jackson, P.J., Hallett, I.C., 2000. Phys-
by these authors. ical and mechanical changes in strawberry fruit after high carbon dioxide
In summary, the results of this study indicate that even though treatments. Postharvest Biol. Technol. 19, 139–146.
the ideal storage temperature for visual appearance of strawberry Harker, F.R., Redgwell, R.J., Hallett, I.C., Murray, S.H., Carter, G., 1997. Texture
fruit is 0.5 ◦ C, acceptable quality of strawberries may be main- of fresh fruit. Hort. Rev. 20, 121–224.
Heinonen, I.M., Meyer, A.S., Frankel, E.N., 1998. Antioxidant activity of berry
tained at moderate temperatures such as 10 ◦ C for a short term phenolics on human low-density lipoprotein and liposome oxidation. J.
storage period of 4 days. However, the RH must be maintained Agric. Food Chem. 46, 4107–4112.
low enough to avoid the onset of fungal decay. Fruit ripening Heo, H.J., Lee, C.Y., 2005. Strawberry and its anthocyanins reduce oxidative
is delayed at 10 ◦ C compared with 20 ◦ C, and may provide a stress-induced apoptosis in PC12 cells. J. Agric. Food Chem. 53, 1984–1989.
useful balance between visual quality attributes and those asso- Jia, Z., Tang, M., Wu, J., 1999. The determination of flavonoid contents in
mulberry and their scavenging effects on superoxide radicals. Food Chem.
ciated with nutritional status. Red ripe berries were used in these 64, 555–559.
experiments and storage periods were relatively short. We are Kalt, W., Forney, C.F., Martin, A., Prior, R.L., 1999. Antioxidant capacity, vita-
currently investigating the effects of fruit maturity and a wider min C, phenolics, and anthocyanins after fresh storage of small fruits. J.
range of RH over extended storage periods in order to further Agric. Food Chem. 47, 4638–4644.
investigate the effects of storage temperature on quality of straw- Liu, M., Li, X.Q., Weber, C., Lee, C.Y., Brown, J., Liu, R.H., 2002. Antioxi-
dant and antiproliferative activities of raspberries. J. Agric. Food Chem. 51,
berries. 2926–2930.
Liu, R.H., 2003. Health benefits of fruit and vegetables are from additive and syn-
Acknowledgements ergistic combinations of phytochemicals. Am. J. Clin. Nutr. 78, 517S–520S.
Meyers, K.J., Watkins, C.B., Pritts, M.P., Liu, R.H., 2003. Antioxidant
and antiproliferative activities of strawberries. J. Agric. Food Chem. 51,
Youngjae Shin was supported by the Ottogi Food Research 6887–6892.
Center, Korea. Darryl Holliday was a Summer Food Science Mitcham, E.J., 2004. Strawberry. In: Gross, K.C., Wang, C.Y., Saltveit, M.E.
Scholar in the Department of Food Science. We also thank (Eds.), The Commercial Storage of Fruits, Vegetables, and Florist and Nurs-
Karina Polar-Cabrera, a Summer Food Science Scholar, and Kari ery Crops. U.S. Department of Agriculture, Agricultural Research Service,
Beltsville Area, Agriculture Handbook 66 on the Website of the USDA
Getchonis for assistance, and Jun Yang, a graduate student in the (http://www.ba.ars.usda.gov/hb66/index.html).
Department of Food Science for assistance with TOSC analy- Nunes, M.C.N., Brecht, J.K., Morais, A., Sargent, S.A., 1998. Controlling tem-
sis. This research was supported by Cornell Univ. Agricultural perature and water loss to maintain ascorbic acid levels in strawberries during
Experiment Station federal funds, project number NE-1018. postharvest handling. J. Food Sci. 63, 1033–1036.
Osborne, T.S., Bacon, J.A., 1961. Two improved and inexpensive systems for
moisture stabilization in seeks and other tissues. J. Plant Physiol. 36, 309.
References Pritts, M.P., Watkins, C.B., 1998. Harvesting, handling, and transporting fresh
fruit. In: Pritts, M., Handley, D. (Eds.), Strawberry Production Guide. North-
Ayala-Zavala, J.F., Wang, S.Y., Wang, C.Y., Gonzalez-Aguilar, G.A., 2004. east Regional Agricultural Engineering Service, Ithaca, NY, pp. 104–108.
Effect of storage temperatures on antioxidant capacity and aroma compounds Rekika, D., Khanizadeh, S., Deschenes, M., Levasseur, A., Charles, M.T., Tsao,
in strawberry fruit Leben. Wissenschaft Technol.: Food Sci. Technol. 37, R., Yang, R., 2005. Antioxidant capacity and phenolic content of selected
687–695. strawberry genotypes. HortScience 40, 1777–1781.
 Y. Shin et al. / Postharvest Biology and Technology 45 (2007) 349–357
Singleton, V.L., Orthofer, R., Lamuela-Raventos, R.M., 1999. Analysis of total
phenols and other oxidation substrates and antioxidants by means of Folin-
Ciocalteu reagent. Oxidants and Antioxidants Pt A, 152–178.
Smith, S.H., Tate, P.L., Huang, G., Magee, J.B., Meepagala, K.M., Wedge, D.E.,
Larcom, L.L., 2004. Antimutagenic activity of berry extracts. J. Med. Food
7, 450–455.
Sun, J., Chu, Y.F., Wu, X.Z., Liu, R.H., 2002. Antioxidant and anti proliferative
activities of common fruits. J. Agric. Food Chem. 50, 7449–7454.
Terada, M., Watanabe, Y., Kunitomo, M., Hayashi, E., 1978. Differential rapid
analysis of ascorbic-acid and ascorbic-acid 2-sulfate by dinitrophenylhy-
drazine method. Anal. Biochem. 84, 604–608.
Vinson, J.A., Su, X.H., Zubik, L., Bose, P., 2001. Phenol antioxidant quantity
and quality in foods: fruits. J. Agric. Food Chem. 49, 5315–5321.
Wang, H., Cao, G.H., Prior, R.L., 1996. Total antioxidant capacity of fruits. J.
Agric. Food Chem. 44, 701–705.
Wang, H., Cao, G.H., Prior, R.L., 1997. Oxygen radical absorbing capacity of
anthocyanins. J. Agric. Food Chem. 45, 304–309.
Wang, S.Y., Feng, R.T., Lu, Y.J., Bowman, L., Ding, M., 2005. Inhibitory effect
on activator protein-1, nuclear factor-kappaB, and cell transformation by
357
extracts of strawberries (Fragaria × ananassa Duch.). J. Agric. Food Chem.
53, 4187–4193.
Watkins, C.B., Manzano-Mendez, J.E., Nock, J.F., Zhang, J.Z., Maloney, K.E.,
1999. Cultivar variation in response of strawberry fruit to high carbon dioxide
treatments. J. Sci. Food Agric. 79, 886–890.
Werner, R.A., Frenkel, C., 1978. Rapid changes in firmness of peaches as influ-
enced by temperature. HortScience 13, 470–471.
Werner, R.A., Hough, L.F., Frenkel, C., 1978. Rehardening of peach fruit in cold
storage. J. Am. Soc. Hort. Sci. 103, 90–91.
Winston, G.W., Regoli, F., Dugas, A.J., Fong, J.H., Blanchard, K.A., 1998.
A rapid gas chromatographic assay for determining oxyradical scavenging
capacity of antioxidants and biological fluids. Free Radic. Biol. Med. 24,
480–493.
Woodward, J.R., 1977. Physical and chemical changes in developing strawberry
fruits. J. Sci. Food Agric. 23, 465–473.
Zhang, J.J., Watkins, C.B., 2005. Fruit quality, fermentation products, and
activities of associated enzymes during elevated CO2 treatment of straw-
berry fruit at high and low temperatures. J. Am. Soc. Hort. Sci. 130, 124–
130.