Maximizing Antioxidants in Fruits

S.Y. Wang
Genetic Improvement of Fruits and Vegetables Laboratory
Plant Sciences Institute, US Department of Agriculture
Beltsville, Maryland 20705-2350
USA

Keywords: antioxidants, free radical, preharvest and postharvest factors, health benefits

Abstract
Fruits contain high levels of antioxidant compounds, such as carotenoids,
flavonoids, vitamins and phenols. These antioxidants are capable of performing a
number of functions including free radical scavengers, peroxide decomposers,
singlet and triplet oxygen quenchers, enzyme inhibitors, and synergists.
Antioxidants can also delay or prevent the oxidation of lipids or other molecules by
inhibiting the initiation or propagation of oxidizing chain reactions. Preharvest
conditions such as climate, temperature, light intensity, soil type, compost mulching,
fertilization, increasing carbon dioxide concentration in the atmosphere, and
application of naturally occurring compounds, all can affect the antioxidant content
and antioxidant activity of the harvested fruits. Other factors affecting antioxidant
activities including crop genotype variation and maturity, culture practices,
postharvest handling and storage are also discussed. Methods to maximize
antioxidants in fruits such as improving selection criteria among different
horticultural cultivars, improving preharvest conditions and postharvest handling
are presented.

INTRODUCTION
Antioxidants are compounds that can delay or inhibit the oxidation of lipid or
other molecules by inhibiting the initiation or propagation of oxidizing chain reactions
(Velioglu et al., 1998). The antioxidant activity of phenolic compounds are mainly due to
their redox properties, which can play an important role in adsorbing and neutralizing free
radicals, quenching singlet and triplet oxygen or de-composing peroxide (Osawa, 1994).
In general, there are two basic categories of antioxidants, natural and synthetic. Recently,
interest has been increasing considerably in finding naturally occurring antioxidants for
use in foods or medicinal materials to replace synthetic antioxidants which are being
restricted due to their possible carcinogenicity (Ito et al., 1983). Fruits and vegetables
contain high levels of phytochemicals in addition to phenolic compounds, such as
nitrogen compounds, carotenoids, and ascorbic acid (Larson, 1988; Velioglu et al., 1998).
Many of these phytochemicals possess significant antioxidant capacities, which provide
protection against harmful free radicals and have been associated with a number of health
benefits (Ames et al., 1993; Cao et al., 1996; Velioglu et al., 1998; Wang et al., 1996).
Diets rich in fruits and vegetables significantly reduce the incidence and the mortality
rates of cancer, cardiovascular disorders, and other degenerative diseases caused by
oxidative stress (Ames et al., 1993; Cao et al., 1996; Velioglu et al., 1998). Eating fruits
and vegetables also could reduce blood pressure, enhances the immune system, detoxifies
contaminants and pollutants, and reduces inflammation (Ames et al., 1993; Larson, 1988;
Velioglu et al., 1998). Therefore, modification of dietary and lifestyle habits can lead to
significant improvements in human health. This paper summarizes the antioxidant
capacities of fruits and describes the factors affecting these antioxidant activities.

ANTIOXIDANT CAPACITY IN FRUITS
The phytochemicals in plant tissues responsible for antioxidant capacity can
largely be attributed to the phenolics, anthocyanins, carotenoids, and other flavonoid
compounds. Fruits and vegetables contain many of these components. In the typical US

81
Proc. 6
th
International Postharvest Symposium
Eds.: M. Erkan and U. Aksoy
Acta Hort. 877, ISHS 2010
diet, daily intake of anthocyanins is approximately 180-215 mg/day, and represents the
largest group of phenolic compounds in the human diet (Khnau, 1976). Anthocyanins
are glycosides and acylglycosides of anthocyanidins. Some common anthocyanidins have
varying hydroxyl or methyl substitutions in their basic structure, flavylium. Over 250
naturally occurring anthocyanins exist, and are differentiated further by their O-
glycosylation with different sugar substitutes (Francis, 1989). Glucose, rhamnose, xylose,
galactose, arabinose, and fructose are the most common sugars substituted on the aglycon
(anthocyanidin). The common anthocyanins are either 3- or 3, 5-glycosylated. Free
radical scavenging properties of the phenolic hydroxy groups attached to ring structures
are responsible for the strong antioxidant properties of the anthocyanins (Rice-Evans and
Miller, 1996; Wang et al., 1997). The structures of the major phenolics in fruits are listed
in Figure 1. Among the anthocyanins, antioxidant strength in preventing oxidation of
human low-density lipoprotein is as follows: delphinidin > cyanidin > malvidin >
pelargonidin (Satu-Gracia et al., 1997). They help reduce damage caused by free-radical
activity and low-density lipoprotein oxidation, platelet aggregation, and endothelium-
dependent vasodilation of arteries (Heinonen et al., 1998; Rice-Evans and Miller, 1996).
The other important flavonoids, and flavones include apigenin, luteolin, diosmetin
and flavonols which include quercetin, myricetin, kaempferol, and their glycosides. They
occur in nearly all our common fruits and vegetables. These flavonoids have antioxidant
and antitumor properties, applications as antibiotics, antidiarrheal, antiulcer and anti-
inflammatory agents and play an important role in the treatment of diseases such as
hypertension, vascular fragility, allergies, hypercholesterolemia (Heinonen et al., 1998;
Khnau, 1976; Larson, 1988; Rice-Evans and Miller, 1996).
Carotenoids are found in many fruits and vegetables such as carrots, dark leafy
greens, orange and yellow vegetables and orange fruits. They are comprised of a set of
several hundred pigmented, fat soluble antioxidants. The most abundant carotenoids in
fruits and vegetables are -carotene, -carotene, lycopene, lutein, zeazanthin, and -
cryptozanthin. These carotenoids account for more than 90% of the carotenoids present in
the human diet. Combination of fatty foods with carotenoid-rich fruits and vegetables
enhance carotenoids uptake. Recent studies have shown that the bioavailability of
lycopene from tomato increased dramatically by heat treatment in the presence of oil
(Gartner et al., 1997). Lycopene was found to be more bioavailable from tomato paste
than from fresh tomato due to heat treatment and the presence of higher oil content in the
paste (Stahl and Sies, 1992).
Recent studies have suggested that -carotene itself may reduce the risk of cancer
and heart diseases even before being converted into vitamin A (Giovannucci, 1999). As
an excellent antioxidant and radical trapping agent, -carotene has the capability to
protect membranes, DNA, and other cellular constituents from oxidative damage (Byers,
1992). Among all carotenoids, lycopene appears to have the greatest anti-oxidant activity,
followed by -carotene, -carotene, lutein, and cryptoxanthin. The singlet oxygen
quenching ability of lycopene is twice that of -carotene and 10 times that of -
tocopherol (Di Mascio et al., 1989; Liebler, 1993).
The antioxidant defense system is composed of different antioxidant components.
Some antioxidant activities in fruits and vegetables may be attributable to other
unidentified substances or to synergistic interactions. There was a positive linear
correlation between the phenolic content and antioxidant capacity of fruits and
vegetables. Therefore, the total antioxidant potential of fruits and vegetables is more
important than individual levels of specific antioxidants (Miura and Nakatani, 1989).
Thus, supplementing fruits and vegetables with a balanced diet is more effective than
consuming high doses of an individual antioxidant such as ascorbic acid, vitamin A or E,
in protecting the body against various oxidative stresses (Wang et al., 1996). This paper
summarizes the antioxidant capacities of various fruit crops and the pre-harvest and
postharvest on affect the antioxidant content and antioxidant activity of the fruits. The
quantity and quality of antioxidants present in fruits could be improved through the
selection of different fruit cultivars and improvement of pre-harvest conditions and

82
postharvest handling techniques.

EFFECT OF PRE-HARVEST FACTORS ON ANTIOXIDANT ACTIVITY OF
FRUITS

Genotype Variation
Genetic factors play an important role in determining antioxidant capacity in
crops. The influence of variety on the content of phenolic compounds and antioxidant
activity in the fruits have been studied by several researchers (Wang and Jiao, 2000;
Wang and Lin, 2000; Howell et al., 2001; Kalt et al., 2001; Siriwoharn et al., 2004;
Taruscio et al., 2004). In berry fruits, Meyers et al. (2003) measured the antioxidant
activities of 8 strawberry genotypes (Annapolis, Allstar, Earliglow, Evangeline,
Mesabi, Sable, Sparkle and Jewel) and found Earliglow (134.13.0 mol vitamin
C/g fw) had the highest total antioxidant activity. The lowest total antioxidant activity
occurred in Allstar, while activity in the other cultivars ranged from 59.9 for Mesabi to
36.7 mol vitamin C/g fw for Jewel. Wang and Lewers (2006) evaluated 19 accessions
from each of the three species, cultivated strawberry plus the two progenitor species (F.
ananassa, F. virginiana and F. chiloensis) and found that fruit of the 13 accessions from
the wild progenitor species, F. virginiana, had significantly higher antioxidant capacity,
total phenolics, and total anthocyanins than did the fruit of three accessions tested from
the cultivated strawberry, F.ananassa, or the three accessions from the other wild
progenitor species, F. chiloensis. Of the evaluated F. virginiana accessions NC 95-19-1,
JP 95-1-1, CFRA 0982, NC 96-5-3, and RH 30 fruit were highest in antioxidant
capacity and should prove useful towards development of strawberry cultivars with high
antioxidant capacities.
Blueberries have a high antioxidant capacity compared with other fruits. Among
the blueberry cultivars, antioxidant activity ranged from 8.1 to 38.3 mol TE/g fw
(Sellapan et al., 2002). Prior et al. (1998) reported an overall range in antioxidant activity
from 13.9 to 45.9 mol TE/g fw in their study of northern and southern highbush,
rabbiteye, and lowbush blueberry genotypes harvested in a single year, with considerable
overlap in antioxidant activity values among the genotypes of different species. The
Premier cultivar from rabbiteye blueberries gave the highest antioxidant capacity value
of 38.29 mol TE/g fw. Climax gave the lowest antioxidant capacity of 19.73 mol
TE/g fw among the rabbiteye blueberries. Howell et al. (2001) found lowbush blueberries
were consistently higher in anthocyanins, total phenolics and antioxidant capacity,
compared with highbush blueberries. Other studies reported that ORAC values among
blueberry genotypes varied considerably (Kalt et al., 1999; Connor et al., 2002;
Ehlenfeldt and Prior, 2001). These data indicate that ample genetic variation exists for
exploitation by plant breeders.
Blackberries and raspberries are also excellent sources of natural antioxidants.
Thomas et al. (2005) studied six cultivars of blackberries and showed Loch Ness had the
highest and Arapaho had the lowest total antioxidant activity. Siriwoharn et al. (2004)
evaluated 11 blackberry cultivars and found ORAC values of blackberries ranging from
37.6 to 75.5 mol TE/g fw and ferric reducing antioxidant power (FRAP) values ranging
from 63.5 to 91.5 mol TE/g fw. ORUS 1489-1, Marion and Evergreen were
varieties containing the highest FRAP values. Deighton et al. (2000) analyzed the
antioxidant properties of 18 domesticated and wild Rubus species and found the
antioxidant capacities of fruit ranged from 0 to 25.3 mol TE/g fw or from 190 to
66,000 mol/L FRAP. Prior et al. (1998) reported R. caucasicus had the highest phenolic
content and antioxidant capacity close to that reported for blueberries.

Maturity
Antioxidant capacity also varies considerably with different stages of maturity.
Many phytonutrients are synthesized in parallel with the overall development and
maturation (Prior et al., 1998; Wang and Lin, 2000; Lester and Eischen, 1996).

83
Strawberry fruits harvested during their ripe stage consistently had higher antioxidant
values than harvested in the pink stage. Beyond the 50% red stage, the antioxidant value
steadily increased with fruit maturity (Wang and Lin, 2000). Immature blueberries
harvested at an early stage (immediately after turning blue) have lower ORAC values and
total anthocyanin content when compared with more mature blueberries harvested 49
days later (Prior et al., 1998). Kalt et al. (2003) found anthocyanin content was
substantially higher in highbush blueberries of more advanced stages of ripeness than in
berries of less ripeness. In grapes, the content of all polyphenolics generally increases as
fruit ripen the highest concentrations being found in the skins (Lee and Talcott, 2004).
Free ellagic acid, ellagic acid glycosides, and total ellagic acid increases during
maturation in the skin of grapes (Lee and Talcott, 2004). Antioxidant activities also
increase with ripening in Rubus L. hybrids (Siriwoharn et al., 2004). Ripening also results
in an increase of total antioxidant activity and vitamin C content in both peel and flesh of
yuzu citrus (Yoo et al., 2004). Beekwilder et al. (2005) found that the dominant
antioxidants in raspberries were anthocyanins, ellagitannins, and proanthocyanidin-like
tannins. Cyanidin 3-glucoside was present in unripe and ripe fruit, while cyanidin
sophoroside, cyanidin glucosylrutinoside, and pelagonidin glucosylrutinoside were
present only when the red color of the fruit developed. The findings presented here
suggest that the content of individual health-promoting compounds in fruits could vary
significantly according to their developmental stage.

Environmental Growing Condition
Preharvest conditions of fruits and vegetables such as growing temperature and
light intensity can affect their antioxidant activities and flavonoid content. Light has been
shown to be an important environmental factor influencing anthocyanin biosynthesis in
plants. Increased light intensity resulted in higher ascorbic acid content in strawberry fruit
compared with the same varieties produced under lower light intensity (Austin et al.,
1960; Wang et al., 2009). In submerged-harvested cranberries, red light and far-red light
increased the total anthocyanin level (Zhou and Singh, 2004). Natural light conditions
enhanced the concentrations of cyanidin 3-galactoside, cyanidin 3-arabinoside, peonidin
3-galactoside, peonidin 3-glucoside, peonidin 3- arabinoside and cyanidin 3-glucoside
substantially (54 to 100%), as compared with the control berries which were kept in the
dark. Red and far-red light had the most prominent effects on cyanidin 3-glucoside and
peonidin 3-arabinoside, showing a 70-92% increase (Zhou and Singh, 2004). Strawberry
growth under high temperature significantly enhanced content of flavonoids and the
antioxidant capacities and plants grown in cool day and night temperature generally had
the lowest antioxidant capacity in fruit (Wang and Zheng, 2001). Month-to-month
variability in vitamin C content depending on growing conditions has been documented in
processed Florida citrus products such as orange and grape-fruit juices (Lee and Coates,
1997). Growing environments have been shown to play a significant role in the
composition of flavonols in red raspberry juice (Rommel and Wrolstad, 1993). These data
suggest a significant influence of climate conditions on flavonoid contents and
antioxidant activity.

Cultural Practices
Different cultural systems have varying effects on antioxidant activity,
anthocyanin, and total phenolic content in various types of fruits. Strawberries from a hill
plasticulture (HC) production system have higher flavonoid content and antioxidant
capacities compared with fruit from plants grown in a matted row (MR) system (Wang et
al., 2002). In general, phenolic acid and flavonol content, as well as cyanidin-and
pelargonidin-based anthocyanins and total flavonoids are greatest in the HC system.
Fruits from plants grown in the MR system generally have the lowest content of phenolic
acids, flavonols, and anthocyanins; but fruit grown under HC conditions have the highest
peroxyl radical absorbance capacity (Wang et al., 2002). Additional compost applied to
the soil also enhanced oxygen absorbance capacity for ROO

, O
2
-
, H
2
O
2
, OH

, and
1
O
2


84
radicals in strawberries (Wang and Lin, 2003). Carbonaro et al. (2002) found a higher
polyphenol content and polyphenoloxidase activity in organic peaches and pears
compared with those of corresponding conventional samples. Ascorbic acid and -
tocopherol were also higher in organic than conventional peaches and pears. Lombardi-
Boccia et al. (2004) found that ascorbic acid, -, -tocopherols, -carotene, myrecitin and
kaempferol were higher in organic plums grown on soil covered with natural meadow,
but total polyphenols and quercetin were higher in conventional plums. Wang et al.
(2008) also showed that ORAC, total anthocyanins, and total phenolic content of
blueberries in organic culture were higher than fruit from the conventional culture. The
organic culture also produced fruit with high chlorogenic acid and flavonoid content.
Thus, antioxidant content in fruits is affected by cultural system.
Plant grown in low-organic-matter and low-cation-exchange-capacity sandy soil
amended with calcium, magnesium, and nitrogen produced more ascorbic acid in their
fruit than plants without supplemental fertilizer (Penalosa et al., 1994). In cantaloupe
fruits, -carotene content was higher in plants grown on silty clay loam soils compared to
plants grown on sandy loam soils (Lester and Eischen, 1996). Carbon dioxide
concentrations in the atmosphere may also have an effect on antioxidant capacity. Higher
CO
2
concentrations in the field resulted in an increase of anthocyanins, phenolics and
antioxidants in strawberry fruit. Strawberries grown in CO
2
-enriched conditions had
higher scavenging capacity for reactive oxygen species and oxygen radical absorbance
activity against ROO

, O
2
-
, H
2
O
2
,

OH, and
1
O
2
radicals than fruit grown under normal
environment without CO
2
enrichment (Wang et al., 2003).

EFFECT OF POST-HARVEST FACTORS ON ANTIOXIDANT ACTIVITY OF
FRUITS

Storage Conditions
Postharvest handling can also affect antioxidant capacity and phytonutrient levels
in fruits depending upon type of fruit, temperature, and storage environment. Increases in
anthocyanin content during storage have been reported for various fruits. Cranberries
stored at temperatures greater than 0C increased antioxidant capacity, anthocyanins and
total phenolic content (Wang and Stretch, 2001). Strawberries stored at 4C showed good
retention of vitamin C, but not chlorgenic acid and quercetin. Both temperature and light
affected the rate of color development in raspberries (Wang et al., 2008). Strawberries
and raspberries stored at temperatures higher than 0C also resulted in an increase in
antioxidant capacity, anthocyanins, and total phenolic content and the magnitude of this
increase was related to storage temperature (Kalt et al., 1999). Ayala-Zavala et al. (2004)
also found that storage temperatures significantly affected the ORAC of strawberry fruit.
Chandler strawberries stored at 10 or 5C had higher antioxidant capacity, total
phenolics, and anthocyanins than those stored at 0C. In general, as storage temperatures
increased, ORAC values also increased. The rate of anthocyanin accumulation increased
with increasing temperature, while flavonols (quercetin and kaempferol derivatives),
ellagic acid and total phenolic contents remained little affected by storage temperatures.
During storage at all temperatures there was a significant increase in the ratio
between pelargonidin and cyanidin for the three varieties of strawberries, showing a
preferential synthesis of pelargonidin over cyanidin (Cordenunsi et al., 2005). Connor et
al. (2002) demonstrated that changes in antioxidant activity, total phenolic, and
anthocyanin content in blueberry during cold storage were variety-dependent. Tarozzi et
al. (2004) found total phenolics

and total antioxidant activity of apple skin but

not in pulp
decreased after 3-6

months of 0C storage. Sluis et al. (2001) showed after 25 weeks of
cold storage, there was no decrease in chlorogenic acid in Jonagold, Golden Delicious,
Red Delicious, Elstar, and Coxs Orange apples, but catechin content decreased
slightly in Golden Delicious, Elstar, and Coxs Orange apples. Goulding et al. (2001)
found little effect on phenolic content in the peel of Granny Smith, Lady Williams,
and Crofton apple stored in air or at 0C for 9 months. However, Leja et al. (2003)

85
showed total phenols, total antioxidant activity, and radical scavenging activity increased
considerably in the peel of two apple cultivars (Jonagold and Sampion) after stored
for 120 days at 1C. Lattanzio et al. (2001) also found that after 60 days of cold storage
the concentration of total phenolics in the skin of Golden Delicious apples increased,
but after 100 days, the total phenolics in the skin began to decrease.

Control Atmospheres
1. High Carbon Dioxide Treatment. High CO
2
atmosphere has been used to retard the
microbial growth and to suppress the respiration rate, ethylene production, softening, and
other metabolic activities associated with the senescence of fresh fruits and vegetables.
The potential benefit from using controlled or modified atmosphere storage is dependent
upon commodity, variety, physiological age, atmosphere composition, and temperature
and duration of storage. Carbon dioxide-enriched atmospheres (10-20%) are especially
effective in retarding decay and softening of strawberries (Gil et al., 1997). However,
exposure to high concentration of CO
2
could adversely affect the color change in
strawberry fruit. High CO
2
caused a reduction in red color intensity and a decrease in
anthocyanin content of internal tissue of strawberry fruit. As CO
2
levels increased, the
concentration of pelargonidin glycosides in the internal tissue decreased (Gil et al., 1997).
Agar et al. (1997) found a decrease in vitamin C content in strawberries associated with
10-30% CO
2
storage. Holcroft and Kader (1999) reported that the concentrations of
ellagic acid, catechin, quercetin and kaempferol derivatives in strawberries increased
during air storage but remained constant during high CO
2
storage.
2. High Oxygen Treatment. Zheng et al. (2003) studied the effects of superatmospheric
O
2
treatments (40, 60, 80, or 100% O
2
at 5C) on Duke highbush blueberries and
showed that antioxidant levels were markedly increased by 60-100% O
2
treatments as
compared with 40% O
2
treatment or air control during 35 days of storage. Elevated O
2

between 60 and 100% also promoted increases in total phenolics and total anthocyanins,
especially malvidin-based anthocyanins as well as the individual phenolic compounds
(Zheng et al., 2003, 2006). High O
2
(40% at 5C) storage of strawberries showed higher
antioxidant capacity, total phenolics, less decay and longer shelf life (Ayala-Zavala et al.,
2007). In comparison with fruits stored in air, strawberries held in 80% O
2
+ 20% CO
2

had higher levels of total anthocyanins during the first 4 days of storage at 8C but
significantly lower levels at the end of 9-day storage (Prez and Sanz, 2001). Chinese
bayberry fruit stored at 5C and 100% O
2
also enhanced antioxidant enzyme activities and
high antioxidant capacities (Yang et al., 2005).
3. Heat Treatment. Heat treatment is used to reduce postharvest quality loss and
suppress pathogen development. However, heat treatment (48C air, 3h) reduced
anthocyanin accumulation and phenylalanine ammonialyase (PAL) activity in
strawberries (Civello et al., 1997). Yoshikawa et al. (1992) treated Chandler
strawberries with humid air at 43 and 46C

for 80 min and found severe damage in the
fruits. Lemons treated with hot-water dip (53C, 3 min) and 1.5% CaCl
2
(15C, -33 KPa,
10 min) reduced chilling injury and maintained antioxidant enzyme catalase and
superoxide dismutase activities (Safizadeh et al., 2007). These contradictory results could
be due to species- and variety-dependent responses of fruits to heat treatments.
4. Illumination (Light and UV) and Ozone (O
3
). Anthocyanins of fruits can be
improved after harvest using artificial light illumination (Austin et al., 1960; Wang et al.,
2009). A range of wavelengths from UV-B (280-320 nm) to red light (680-780 nm) is
effective. Red light alone is only slightly effective in stimulating anthocyanin production,
UV-B has an additional effect, and both together have synergistic effects. UV treatment
enhanced anthocyanin levels in strawberries (Baka et al., 1999). Pan et al. (2004) showed
that UV-C (4.1 kJ m
-2
) and heat treatment (45C), either separately or combined, all
reduced anthocyanins and phenolics in Seascape strawberries compared with controls.
However, UV-C (9.2 kJ m
-2
) and heat treatments (45C) retained fruit quality and
antioxidant activity better than in control fruit of boysenberries (Vicente et al., 2004).

Wang et al. (2009) found higher antioxidant capacity was detected in blueberries treated

86
with 2.15, 4.30, or 6.45 kJ m
-2
compared with control fruit. UV-C dosage of 0.43 kJ m
-2

also increases phenolics and anthocyanins, but to a lesser extent. The optimum doses of
UV-C for enhancing phytochemical content in blueberries were 2.15 and 4.30 kJ m
-2
. The
enrichment of stilbenes, such as resveratrol, in grapes in response to stress induced by
postharvest UV-C irradiation has been described (Cantos et al., 2002). The increases of
total phenols and anthocyanins in blueberries by UV-C illumination appeared to be dose-
dependent at lower doses (0.43~2.15 kJ m
2
). However, higher doses (4.30~6.45 kJ m
2
)
tended to suppress these increases. This phenomenon has also been reported in
strawberries where high doses of UV-C exposure was thought to cause too much stress
and possibly resulted in injury (Baka et al., 1999). These results indicate that adequate
UV irradiation of blueberries can be beneficial in terms of increasing the levels of
antioxidant activity and potentially health-promoting nutraceutical compounds.
Ozone (O
3
) is an unstable compound which produces hydroxyl radicals and other
free radical species. Ozone has been used in different applications in the food industry,
and has been recommended as a GRAS (generally recognized as safe) disinfectant or
sanitizer for foods in the United States (Graham, 1997). Barth et al. (1995) and Perez et
al. (1999) reported that blackberries and strawberries stored in an atmosphere containing
ozone atmosphere (0.3-0.35 l/L) showed a sharp decrease in anthocyanin levels.

Treatment of Natural Compounds or Chemicals
1. Methyl Jasmonate (MJ), 1-Methylcyclopropene (1-MCP) and Allyl Thiocyanate
(AITC). Methyl jasmonate (MJ) (Fig. 2), a naturally occurring compound, was found to
maintain higher levels of oxygen radical absorbance capacity for ORAC, O
2
-
, H
2
O
2
,

OH,
1
O
2,
DPPH* and ABTS
+
in cranberries, strawberries, raspberries, blueberries and
blackberries (Changjirakul et al., 2006, 2007). Ayala-Zavala et al. (2005)

found that
Allstar strawberry fruit treated with MJ in conjunction with ethanol showed higher
antioxidant capacity, total phenolics, and anthocyanins than those treated with ethanol
alone or controls (non-treated) during the post-harvest period.
1-MCP has been used to extend postharvest life of many fruits. Red Delicious
apple treated with 1-MCP improves the retention of flavonoids (MacLean et al., 2006).
Yali pear treated with 1-MCP also increases activities of antioxidant enzymes (catalse,
superoxide dismutase, and peroxidase), antioxidant potential and improves postharvest
quality (Fu et al., 2007). Japanese plum treated with 1-MCP reduces ethylene production,
softening and extends the storage life, but antioxidants showed no effect (Khan and
Singh, 2008). Strawberry (Everest) fruit treated with 1-MCP inhibited PAL activity and
slowed increases in anthocyanin and phenolic content (Jiang, 2001).
AITC is a natural compound which is present in all plants belonging to the
Cruciferae family and is generally considered safe for human consumption and believed
to be conducive to health (Kermanshai et al., 2001; Shin et al., 2004; Troncoso et al.,
2005). Although it is not considered to be a nutrient in the classical sense, AITC is
bioactive and has been shown to have various biological effects including anti-oxidative,
anti-bacterial, anti-fungal, anti-nematode and anti-insect activities (Kermanshai et al.,
2001; Shin et al., 2004). It has been shown that blue mold (Penicillium expansum) in
pears was controlled by AITC vapor treatment (Mari et al., 2002). Furthermore, the
antimicrobial activity of AITC against pathogens in iceberg lettuce, apples, and tomatoes
has also been reported (Lin et al., 2000). Chanjirakul et al. (2006, 2007) have also shown
that AITC reduced decay of strawberries, blackberries and raspberries.
However, the mechanism of AITC in reducing microbial growth is different from
MJ, 1-MCP and essential oils. AITC does not increase antioxidants or antioxidant enzyme
activity, but promoted the accumulation of H
2
O
2
radicals to inhibit the growth and
proliferation of microbial cells, thereby reducing decay in fruit tissue (Wang et al.,
unpublished).
2. Essential Oils or Natural Volatile Compounds. Essential oils or natural volatile
compounds are aromatic oily extracts obtained from plant materials such as buds, flowers,
seeds, leaves, barks, roots, fruits, and other parts of the plants such as thymol, menthol,

87
eugenol, carvacrol, anethole, cinnamaldehyde, cinnamic acid, perillaldehyde, linalool, and
p-cymene. Essential oils have also been demonstrated to be effective in inhibiting
microbial growth, reduce fruit decay, enhances phenolic acid and flavonoid content and
free radical scavenging capacity in fruits (Wang et al., 2007, 2008). From electron spin
resonance (ESR) measurement, higher radical scavenging capacities were found against
and 2, 2-Di (4-tert-octylphenyl) -1-picrylhydrazyl (DPPH*) and OH in treated fruit,
particularly in berries treated with thymol, compared to those in control groups (Wang et
al., 2007, 2008). Thus, these natural products have the potential to increase the resistance
of tissues against oxidative damages and to preserve the quality and safety of fresh
produce.

CONCLUSIONS
The level and activity of antioxidants in fruits and vegetables can be affected by
both preharvest and postharvest factors. For example, climate, temperature, cultural
practices and light all affect antioxidant activity in various horticultural crops. The stage
of crop maturity also influences the antioxidant capacity of blueberries, blackberries,
raspberries, strawberries and other fruit and vegetable crops. Postharvest factors such as
pre-storage treatment, and storage temperature and environment can also influence
phytochemical composition of food crops. New research is ongoing to assess the impact
of different handling techniques on the nutritional content of fruits and vegetables. One of
the most significant applications of postharvest technologies relating to nutrition may well
be to retard the softening process of fruits and vegetables so that they can be harvested
and marketed at a more mature stage, when more of the phytonutrients have already been
bio-synthesized. Knowledge gained from these research efforts will be helpful in
maximizing the nutrient content and improving the quality of fruits and vegetables for
human consumption.

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Figures








Fig. 1. The chemical structures of the major phenolics in fruits.












COOH R
1
OH

CH
3
O
HO COOH
COO
HO
HO
HO
HO
OH C
O
OH
R
1
=OH R
2
=H : Cyanidin
R
1
=OH R
2
=OH : Delphinidin
R
1
=OCH
3
R
2
=OH : Petunidin
R
1
=OCH
3
R
2
=OCH
3
: Malvidin
R
1
=OCH
3
R
2
=H : Peonidin
R
1
= H R
2
= OH : Quercetin
R
1
= H R
2
= H : Kaempferol R
1
=
OH R
2
= OH : Myricetin
R
1
=OH : Caffeic acid Vanillic acid Chlorogenic acid
R
1
=H : p-Coumaric acid
R
1
=OCH
3
: Ferulic acid
Ellagic acid
Catechin
Procyanidin Oligomers n=0-13

93







94