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Ascorbic Acid Extends the Shelf-life of Abiu (Pouteria caimito) Fruit by

Maintaining Quality and Delaying Browning Symptoms

Article in The Horticulture Journal · April 2023

DOI: 10.2503/hortj.QH-053

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The Horticulture Journal Preview The Japanese Society for

doi: 10.2503/hortj.QH-053
JSHS Horticultural Science
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Ascorbic Acid Extends the Shelf-life of Abiu (Pouteria caimito) Fruit by

Maintaining Quality and Delaying Browning Symptoms

Slamet Susanto1*, Abdullah Bin Arif2, Siti Mariana Widayanti2 and Deden Derajat Matra1

1
Department of Agronomy and Horticulture, Faculty of Agriculture, IPB University (Bogor Agricultural University), Bogor 16680,
Indonesia
2
Research Center for Agroindustry, National Research and Innovation Agency, Jakarta 10340, Indonesia

Abiu is a tropical fruit with many beneficial bioactive compounds. However, its economic value is limited by
a short shelf life and rapid browning. This study evaluated the effect of ascorbic acid on extending abiu fruit
shelf life during storage. The fruit was soaked in ascorbic acid solutions of 0 mM (control), 5 mM (AA5), and
10 mM (AA10) for 10 minutes, air-dried, then stored in an ambient room (28 ± 1°C and 80 ± 5% RH) for 12
days. The results indicated that the 10 mM exogenous ascorbic acid treatment increased abiu fruit shelf life up
to 12 days, nine days longer than the control. Abiu fruit, after the AA10 treatment, underwent a 1.4-fold lower
weight loss than the control. In addition, on day 12 of storage, the browning of fruit with the AA10 treatment
were 8 and 11% on the peel and pulp, respectively. The climacteric peak of abiu fruit in the AA10 treatment
occurred on day 8 of storage, three days later than the control. The AA10 treatment also maintained vitamin C
content and fruit firmness. Thus, the AA10 treatment effectively extended shelf life and maintained abiu fruit
quality.

Key Words: climacteric, firmness, storage, weight loss.

Introduction
Fruits reduce risks of diseases, including cancers
and cardiovascular disease, support increased fruit
consumption (Isabelle et al., 2010). Abiu fruit is a trop‐
ical fruit that contains important bioactive compounds
such as 1-(2-Hydroxyethyl)-1,2,4-triazole, 5-hydroxy
methyl furfural, 1-methyl-5-fluorouracil, and trans-
geranylgeraniol (Arif et al., 2021), known anti-cancer
agents (Al-Soud et al., 2004; Cristensen et al., 2019;
Kumar and Periyasamy, 2016; Zhao et al., 2013). Addi‐
tionally, the abiu fruit is reported to have anti-microbial
properties (Abreu et al., 2019), inhibit acetyl‐
cholinesterase (Fernandez et al., 2020), and exhibit anti-
cancer activity (Veeramani et al., 2021). However, abiu
fruit suffer from a short shelf life, and rapidly turn
brown (Arif et al., 2022).
The abiu fruit is climacteric, with a steep respiration
peak and high levels of ethylene production, which lim‐
its its shelf life (Arif et al., 2022). The shelf life of har‐
Received; October 21, 2022. Accepted; January 20, 2023.
First Published Online in J-STAGE on April 25, 2023.
Special Issue ‘Postharvest Biology and Technology of Tropical/
Subtropical Horticultural Crops’.
* Corresponding author (E-mail: slmtsanto@gmail.com).
vested fully ripe abiu fruit is roughly six days, and, after
two days of storage, it undergoes a significant increase
in browning index (Arif et al., 2022). These qualities
limit the market distribution of abiu fruit. Furthermore,
peel browning is a major problem for fruit storage and
processing. Browning is reported to be responsible for
more than 50% of fruit industry losses (Jaeger et al.,
2018). Therefore, postharvest treatments are required to
maintain fruit quality and increase shelf life.
Ascorbic acid is an additive that maintains both fruit
quality and extends shelf life. Coating treatments con‐
taining ascorbic acid delayed 1-aminocyclopropane-1-
carboxylic (ACC) production in guava (Martinez-Ortiz
et al., 2019), and diminished weight loss in strawberries
(Sogvar et al., 2016). In addition, pre-harvest ascorbic
acid treatments are reported to inhibit the polygalac‐
turonase (PG) enzyme and maintain grape firmness
(Lo’ay and El-Boray, 2018). Moreover, ascorbic acid is
an antioxidant widely used to prevent fruit browning in
strawberries (Sogvar et al., 2016), mangoes (Lo’ay and
Ameer, 2019), and guava (Azam et al., 2021).
According to Moon et al. (2020), there are three main
mechanisms behind the ascorbic acid inhibition of
browning: 1) ascorbic acid acts as an antioxidant, pro‐
moting o-quinone regeneration and limiting polymer‐
ization into brown pigments; 2) ascorbic acid binds to

© 2023 The Japanese Society for Horticultural Science (JSHS), All rights reserved.
 2 S. Susanto, A. B. Arif, S. M. Widayanti and D. D. Matra
histidine residues which are PPO catalytic by-products,
and reduces the turnover of oxidized phenols induced
by PPO enzymes; 3) ascorbic acid, is a weak acid, and
its accumulation can lower the pH of the cytosol, also
reducing oxidase enzyme activities.
The influence of ascorbic acid on postharvest fruit
quality and abiu fruit browning response during storage
is not yet evaluated. Therefore, this study examines the
effect of ascorbic acid on extending abiu fruit shelf life
during storage in terms of physical properties (brown‐
ing, color, and firmness) as well as chemical properties
including total phenol content (TPC), polyphenol oxi‐
dase (PPO) activity, malondialdehyde (MDA) concen‐
tration, respiration and ethylene production rates, sugar
content and concentration, titratable acidity and vitamin
C content.
Materials and Methods
Fruit materials and storage treatment
Abiu fruit were obtained from farmers' gardens in
West Java Province, Indonesia. Located at longitude
60.21°E, latitude 106.44°S, and 200 m above sea level.
The fruit were harvested when fully ripe (65 days after
fruit set) with bright yellow peel color. Harvest was per‐
formed in the morning, and fruit were brought to the
laboratory within 30 minutes. On arrival, surface dirt
was removed by dipping into water then dried by air
blower. Fruit were then selected for color, shape,
absence physical defects, and weight (250 ± 25 g) to
ensure uniform characteristics. Fruit were then separat‐
ed into treatment groups of 25 fruit in triplicate (75
fruits total per treatment). Each treatment replicate was
soaked in the respective treatment solution for 10 min‐
utes. The treatments were: water (control), 5 mM ascor‐
bic acid (AA5), and 10 mM ascorbic acid (AA10).
After treatment, the fruit were air-dried and placed in a
container (80 cm × 60 cm × 45 cm) without a lid, then
stored in an ambient room (28 ± 1°C and 80 ± 5% RH)
for 12 days. Fruit sampling was carried out on days 0,
3, 6, 9, and 12 (storage periods).
Measurements of weight loss and fruit firmness
Fruit weight loss was calculated using the method
proposed by Kaewjumpol et al. (2021). The difference
between fresh fruit weight before and after storage was
calculated and expressed as a percentage of the initial
fresh weight. The weight loss was expressed in a per‐
centage (%).
The fruit firmness was measured using a Brookfield
digital texture analyzer. The texture analyzer was set to
mode 20, with a maximum load of 10 kg, a pressing
depth of 15 mm, a speed of 60 mm·min−1, and a needle
probe diameter of 2 mm. The firmness measurement of
the abiu fruit was taken in triplicate at the equatorial
section of the fruit. Firmness is expressed in Newton
(N) (Arif et al., 2022).
Peel fruit color and browning index (BI) determination
Color changes were measured according to the
method described by Chinwang et al. (2011). Peel fruit
color was evaluated for lightness (L) and hue angle at 3
different positions using a Colorimeter CR-400 Chroma
Meter (Konica Minolta Inc., Tokyo, Japan). Then, the
average was determined.
The BI was evaluated by the method described by
Arif et al. (2022). Browning was assessed by measuring
the percentage of browning area on fruit peel and pulp,
then assigned a score using the following scale: 0 = no
browning; 1 = < 25% browning; 2 = 25–50% browning;
3 = 50–75% browning; 4 = > 75% browning. The BI
was calculated using the following equation: BI = [Σ
(browning scale) × (number of fruits in each scale)/
(total number of fruits × the highest scale number)] ×
100. The BI was expressed in a percentage (%). The
non-marketability of fruit was indicated by a peel
browning index ≥ 40% (Lichanporn et al., 2020).
Measurements of total phenolic content (TPC),
Polyphenol Oxidase (PPO), and Malondialdehyde
(MDA)
The TPC in fruit pulp was analyzed by the Folin-
Ciocalteu method (Chimvaree et al., 2020). Whereby
150 μL of supernatant, 2400 μL of distilled water, and
150 μL of 0.25 N Folin-Ciocalteu were thoroughly
mixed in a test tube by vortex. The mixture was
allowed to react at room temperature for 3 min, and
300 μL of 1 N Na2CO3 solution was added. The solu‐
tion was well mixed and incubated at room temperature
(25°C) for 2 h. Then, a 200 mL sample, standard or
blank from the assay tube was transferred to a clear 96-
well microplate and the absorbance at 765 nm recorded
for each well. TPC was then evaluated based on
absorbance at 765 nm. The standard gallic acid (GAE)
curve was plotted, with TPC concentration expressed as
mg GAE·kg−1 fresh weight.
The PPO activity was measured according to the
method described by Jiang (1999) with some modifica‐
tions. The sample (1 g) was homogenized in 4 mL
0.1 M sodium phosphate buffer (at pH 7.2) and cen‐
trifuged at 12,000 × g for 30 min at 4°C. The super‐
natant was collected as an enzyme extraction solution.
Then 100 mmol·L−1 citrate buffer (1.45 mL, pH 6.8)
was added to 50 μL of the crude enzyme extract and
0.50 mL of 4-methyl catechol solution (100 mmol·L−1).
One PPO activity unit (U) was defined as the quantity
of enzyme that increased the absorbance at 420 nm by
0.001 per minute. PPO activity is expressed as U·g−1
protein.
The MDA concentrations were measured using the
method of Xi et al. (2017). The sample (1 g) was
homogenized in 5.0 mL of 5% (w/v) trichloroacetic
acid (TCA) and centrifuged for 20 min at 10,000 × g.
Next, the supernatant was collected and mixed with
0.67% TBA in a 1:1 (v/v) ratio, heated at 100°C for
 Hort. J. Preview
20 minutes, and immediately cooled on ice. Then cen‐
trifuged at 10,000 × g for 10 min, with the absorbance
of the supernatant at 450, 532, and 600 nm measure by
spectrophotometer (UV-2600; Shimadzu Co., Kyoto,
Japan). Finally, the MDA concentration was calculated
using the formula: 6.45 × (A532 − A600) − 0.56 × A450.
The MDA concentration was expressed as μmol·kg−1
based on fresh weight.
Respiration, ACC, and ethylene production
The respiration rate and ethylene production during
storage were measured using a Gas Analyzer (Type
F-950; Felix instruments, Camas, WA, USA) by flow‐
ing air in a sealed glass bottle into the Gas Analyzer
with an airflow rate of 70 mL·min−1. The respiration
rate is expressed as mL·kg−1·h−1, and ethylene produc‐
tion is expressed as μL·kg−1·h−1 (Arif et al., 2022).
The ACC quantification was measured according to
Lizada and Yang (1979) method. The GC Merck Varian
450 was used with a detector temperature of 100°C and
a column temperature of 80°C with a gas flow rate of
N2 0.6 mL·min−1. The ACC quantification is expressed
as mL·kg−1.
Soluble sugar concentration and content (sucrose, glu‐
cose, and fructose)
The soluble sugar concentration (SSC) of the fruit
juice was determined using a digital Atago PR-1 refrac‐
tometer (Atago Co. Ltd., Tokyo, Japan) at 28 ± 1°C
according to the method of Asiche et al. (2017) and
expressed as Brix (°Brix).
The sugar content (glucose, sucrose, and fructose)
was measured following the method reported by Zhou
et al. (2020). Aquabidest was added abiu pulp (5 g)
until the volume reached 20 ml, ultrasonically agitated
for 30 min, then passed through a microfilter. Super‐
natants were analyzed using ultra-high performance liq‐
uid chromatography (UHPLC) Merk Dionex Ultimate
3000 RS Column Compartment (mobile phase: acetoni‐
trile, column: C18, flow rate: 1 mL·min−1, detector:
change aerosol detector, and temperature: 25°C). The
sucrose, glucose, and fructose contents are expressed as
percentages.
Titratable acidity and vitamin C
The NaOH titration method described by Asiche
Weight loss (%)
et al. (2017) was used to determine the titratable acidity
(TA) content. TA was determined by titrating the
extract against 0.1 N NaOH and then expressed as per‐
centage citric acid equivalents.
Vitamin C content was evaluated using the method of
Klimczak and Swiglo (2015). Abiu pulp (1 g) was
placed into a scaled test tube, with 0.0065 N H2SO4
added until the volume reached 20 mL, ultrasonically
agitated for 30 minutes, then filtered through a 0.22 μm
filter membrane. Vitamin C content was analyzed by
UHPLC Merk Dionex Ultimate 3000 RS Column
3
Compartment (mobile phase: H2SO4; column: variant
C18, 150 mm long, 4.6 mm in diameter, and 5 μm parti‐
cle size; flow rate: 1 mL·min−1, detector: diode array
detector; wavelength: 254 nm; and temperature: 25°C).
Vitamin C was expressed as mg·10−2·g−1.
Statistical analysis
Experiments were performed using a randomized
complete block design with three replications. Means
were compared by one-way analysis of variance and
Duncan multiple range test (DMRT) at a 5% signifi‐
cance level. Data are presented as mean ± standard
error (SE).
Results and Discussion
Weight loss and fruit firmness
Weight loss is critical in determining fruit’s potential
shelf life (Ribeiro and de Freitas, 2020). Weight loss
may occur due to increased metabolic activity and/or
decreased membrane integrity caused by increased
postharvest senescence (Koyuncu et al., 2019). This
current study observed continual weight loss in abiu
fruit during storage, but weight loss for abiu fruit with
the AA10 treatment was less than the control (Fig. 1).
Weight loss in the abiu fruit on day 12 of storage
reached 24.25, 24.12, and 17.74% for the control, AA5
treated, and AA10 treated fruit, respectively (Fig. 1).
This demonstrated that the AA10 treatment reduced
fruit weight loss by 6.51%, a 27% improvement on the
control. Exogenous application of ascorbic acid to
lychee fruit inhibited postharvest senescence and
weight loss (Ali et al., 2021) and in strawberries,
reduced water loss and weight loss during storage
(Sogvar et al., 2016). Thus, a 10 mM ascorbic acid
treatment will likely reduce metabolic activity, delay
senescence, and maintain cellular integrity, thereby
reducing weight loss in ambiently stored abiu fruit.
Fruit firmness is an important parameter consumers
use in assessing fruit quality. The process of fruit ripen‐
ing induces cell wall modifications, decreasing fruit
firmness (Wang et al., 2018; Zhang et al., 2019). As
30
AA10
AA5 a
control a
20
b
a b
b
10 b
a
0
3 6 9 12
Days after storage
Fig. 1. Effect of ascorbic acid treatments on abiu fruit weight loss
during storage. Data are means of three replicates ± SE. Differ‐
ent letters in the same day indicate significant difference by the
Duncan multiple range test (P < 0.05).
 4 S. Susanto, A. B. Arif, S. M. Widayanti and D. D. Matra

12 94
a
a a a a AA10 AA5 control
b a a
b b
c b 88 a
a
8 c a a a
Firmness (N)

a
b a
AA10 a

Hue
82 b
AA5 b
b
4 control
76

0 70
0 3 6 9 12 0 3 6 9 12
Days after storage Days after storage
Fig. 2. Effect of ascorbic acid treatments on abiu fruit firmness dur‐
ing storage. Data are means of three replicates ± SE. Different (a)
letters in the same day indicate significant difference by the 80 a
Duncan multiple range test (P < 0.05). a a a a
b a a
a
60 b
b b

Lightness
shown in Figure 2, abiu fruit firmness decreased gradu‐
40
ally for all treatments during storage. On day 12, the
AA10
control fruit firmness was 8.05 N. However, the abiu
AA5
fruit firmness with AA10 treatment was 9.87 N, 1.23- 20
control
fold firmer than the control and 1.10-fold firmer than
the AA5 treatment (Fig. 2). In this study, the higher 0
concentration exogenous ascorbic acid treatment main‐ 0 3 6 9 12
Days after storage
tained abiu fruit firmness better during storage. In
grapes, ascorbic acid is known to inhibit polygalac‐ (b)
turonase (PG) enzyme activity, and through this,
Fig. 3. Effect of ascorbic acid treatments on color; a) hue, and b)
maintain grape firmness (Lo’ay and El-Boray, 2018). lightness of abiu fruit during storage. Data are means of three
Additionally, ascorbic acid may maintain fruit firmness replicates ± SE. Different letters in the same day indicate
by reducing membrane lipid peroxidation, enhancing significant difference by the Duncan multiple range test
cell capacity to scavenge ROS, and decreasing fruit (P < 0.05).
respiration (Liu et al., 2014).

Color to tissue darkening (Peng et al., 2021). Thus, the AA10

Color is an essential external indicator of fruit quali‐
ty. Hue and lightness values in abiu peel decreased for
all treatments during storage (Fig. 3). The average hue
and lightness of the abiu peel at harvest were 87.99 and
72.71, respectively. During storage, the AA10 treatment
inhibited the decrease in hue and lightness values of
abiu fruit. At the end of storage, the hue value of the
AA10 and AA5-treated fruit were higher than the con‐
trol (Fig. 3a). This behavior may be due to ascorbic acid
delaying fruit color development (Martinez-Ortiz et al.,
2017). In addition, ascorbic acid can also suppress eth‐
ylene production. This is important as ethylene is essen‐
tial in fruit pigmentation to regulate the expression of
enzymes that influence fruit color (Montalvo et al.,
2009). Abiu peel exhibited similar changes, with day 12
lightness decreased compared to fruit at day 0 of stor‐
age by 19.02, 7.64, and 2.85 units, for control, AA5-,
and AA10-treated fruit, respectively (Fig. 3b). The L
value in abiu fruit also relates to the progress of brown‐
ing during storage. The L value correlated negatively
with litchi browning index, with decreased L values
indicated a higher browning percentage (Zhang et al.,
2015). Discoloration during browning primarily relates
treatment of abiu fruit delays decreases in hue and L
values during storage.
Browning index (BI)
Browning is a primary factor dictating shelf life and
fruit quality during storage (Li et al., 2021; Wang et al.,
2021). Abiu fruit peel and pulp browning gradually
increased during storage (Fig. 4). An important conse‐
quence of browning is the negative impact on consumer
preference. In apples, severe browning (> 30% relative
area) is reported to result in 50% of consumers discard‐
ing the whole apple (Jaeger et al., 2018). Lychee fruit
also loses marketability when surface browning exceeds
a score of 3.0 (Ali et al., 2021). Browning of the abiu
peel increased gradually during storage for all treat‐
ments (Fig. 5a). However, peel browning in the control
was higher than for ascorbic acid-treated fruit during
storage. Overall, compared to the control, the ascorbic
acid treatment inhibited abiu peel browning by 6.25-
fold and 1.88-fold for the AA10 and AA5 treatments,
respectively. The BI in abiu peel, on days 3 and 6 of
storage was > 10% for both the control and AA5 treat‐
ment, while the AA10 treatment BI remained < 8%
 Hort. J. Preview 5

until day 12 of storage.
In addition, the pulp browning index of the control
was > 50% by day 12 of storage (Fig. 5b). On day 12,
pulp browning of the AA10- and AA5-treated fruit was
inhibited by 5-fold and 1.47-fold, respectively, com‐
pared with the control. For the control and AA5-treated
fruit, a pulp BI > 30% occurred by day 3 and day 6,
respectively. For the AA10-treated fruit, the pulp BI
remained < 11% until day 12 of storage. Therefore, the
control abiu fruit lost market value after three days of
storage, while the AA10-treated abiu fruit retained mar‐
ketable value until day 12 of storage. Thus, the AA10
treatment effectively extends shelf-life and given the
simple treatment process, appears appropriate for use in
the abiu fruit industry.
As an antioxidant, ascorbic acid promotes o-quinone
regeneration and prevents polymerization into brown
pigments (Franck et al., 2007; Moon et al., 2020).
Antioxidants react with oxygen, thus inhibiting the ini‐
tiation of browning. Despite ascorbic acid not directly
interacting with PPO enzymes, it inhibits enzymatic
browning by chemically reducing oxidized substrates.
This chemical reduction process of ascorbic acid is
reported to be reduction of the enzymatically formed
o-quinone to di-phenol (Moon et al., 2020).

Fig. 4. The browning symptom in abiu fruit during storage. AA5 = TPC, PPO, and MDA
ascorbic acid 5 mM, and AA10 = ascorbic acid 10 mM.
Enzymatic browning is caused by three main factors:
phenolic compounds, polyphenol oxidases, and oxygen
60 (Moon et al., 2020). Phenolic compounds are substrates
AA10 AA5 control a a for the PPO enzymes involved in the browning reaction
(Zhang et al., 2022). TPC in abiu fruit decreased during
Peel Browning index (%)

40 a storage for all treatments (Fig. 6). At harvest, the TPC

b average was 99.85 mg·10−2·g−1; on day 12 of storage,
b
the TPC of abiu fruit in the control dropped 7.68-fold.
a b
20 However, the AA10 treatment inhibited phenolic degra‐
c dation significantly more than the control and AA5
b c
c
treatment. On average, AA10-treated abiu fruit exhibit‐
a
0
c ed 2.11-fold higher concentrations of TPC than the con‐
0 3 6 9 12 trol and 1.38-fold higher than the AA5 treatment on day
Days after storage
12 of storage. Phenolics after acid treatment were
(a) higher than controls in lychee fruit (Ali et al., 2019;
60
Kumari et al., 2015), mango (Lo’ay and Ameer, 2019),
AA10 a
a
AA5 a
Pulp Browning Index (%)

control
b b 120 AA10 AA5 control
40
b a
a
TPC (mg · 10-2 · g -1)

b 80
20 a
c c
b
c a
a c c a
0 40
b a
0 3 6 9 12
b b
Days after storage
c
0
(b) 0 3 6 9 12
Days after storage
Fig. 5. Effect of ascorbic acid treatments on; a) peel browning
index, and b) pulp browning index of abiu fruit during storage. Fig. 6. Effect of ascorbic acid treatments on abiu fruit total pheno‐
Data are means of three replicates ± SE. Different letters in the lic content during storage. Data are means of three replicates ±
same day indicate significant difference by the Duncan multiple SE. Different letters in the same day indicate significant differ‐
range test (P < 0.05). ence by the Duncan multiple range test (P < 0.05).
 6 S. Susanto, A. B. Arif, S. M. Widayanti and D. D. Matra
1.5
AA10
AA5
1.2
control a b
a b
PPO (U ∙ g-1)
0.9
c b
0.6 b a
a c
0.3
0.0
0 3 6 9
Days after storage
Fig. 7. Effect of ascorbic acid treatments on abiu fruit PPO during
storage. Data are means of three replicates ± SE. Different
letters in the same day indicate significant difference by the
Duncan multiple range test (P < 0.05).
and guava (Lo’ay and Taher, 2018). The phenolic con‐
tent of all abiu fruit decreased, associated with
increased enzymatic activity. However, this was more
pronounced for the control fruit than for treated fruit, as
more damaged cells lead to increased enzyme activity
and increased browning. Phenolic compounds are PPO
enzyme substrates, involved in fruit browning reactions.
As an antioxidant, ascorbic acid inhibits phenol degra‐
dation and the conversion of phenol to quinone (Moon
et al., 2020). Ascorbic acid also inhibits the browning
reaction by reducing o-quinones back to the phenolic
substrates (Soliva-Fortuny and Martín-Belloso, 2003).
Thus, the AA10 treatment for abiu fruit suppresses
phenolic compound oxidation, which inhibits further
browning.
Besides phenolic compounds, PPO enzyme activity
is another factor in browning (Liu et al., 2022). PPO
enzyme activity was observed to increase for all treat‐
ments during storage (Fig. 7). PPO enzyme activity was
observed to increase 6-, 5-, and 5-fold during storage
for the control, AA5, and AA10 treatments, respective‐
ly. PPO enzyme activity is optimal at pH 5–7, with a
lower pH inhibiting enzymatic activity (Queiroz et al.,
2008). As a weak acid, ascorbic acid decreases pH,
inhibiting PPO enzyme activity. However, PPO enzyme
activity in ascorbic acid treated abiu fruit increased dur‐
ing storage. This indicates that the ascorbic acid does
not react directly with the enzyme in suppressing PPO
enzyme activity. Indeed, Arias et al. (2007) reported, in
pears, that rather than direct PPO enzyme activity inhi‐
bition by ascorbic acid, it acts on the oxidized substrate
molecules.
MDA is a product of membrane lipid peroxidation
and is a useful indicator for assessing oxidative damage
to plant tissues (Saleem et al., 2021). The MDA con‐
centration in abiu pulp increased linearly from day 0 to
day 12 (Fig. 8). The AA10 treatment of abiu fruit
reduced MDA production more than other treatments.
The MDA concentration of exogenously AA10-treated
Abiu fruit was 1.41-fold lower than the control (Fig. 8).
0.3
AA10 a
a a
a AA5
MDA (mol ∙ kg -1)
b 0.2
control a b
b
b c
a c c
0.1
a
12 0.0
0 3 6 9 12
Days after storage
Fig. 8. Effect of ascorbic acid treatments on abiu fruit MDA during
storage. Data are means of three replicates ± SE. Different
letters in the same day indicate significant difference by the
Duncan multiple range test (P < 0.05).
As an antioxidant, ascorbic acid prevents reactive oxy‐
gen species (ROS) formation and reduces oxidative
stress in tissues (Ali et al., 2021; Saleem et al., 2021).
One ROS product is MDA, with MDA accumulation
damaging cell membranes composition, promoting
brown polymer formation, and increasing browning
(Lin et al., 2016). Increased MDA content damages cel‐
lular membrane structural integrity, promoting contact
between PPO enzymes and phenolic substrates, leading
to additional browning (Lin et al., 2014). Therefore, the
AA10 treatment of abiu fruit may significantly inhibit
increases in MDA content during storage.
Respiration, ACC and ethylene production
Ascorbic acid treatment affected respiration rate,
ACC concentration, and ethylene production (Fig. 9).
The CO2 production of abiu fruit decreased over the
first three days of storage then increased until the cli‐
macteric peak was reached for each treatment (Fig. 9a).
The peak respiration rate (CO2 production and O2 con‐
sumption) occurred for the control fruit on day 5
(Fig. 9a, b), while, for the AA10-treated fruit, the peak
respiration rate occurred on day 8. Ascorbic acid treat‐
ment is previously reported to suppress the respiration
rate in guava (Martinez-Ortiz et al., 2019). In addition,
the O2 concentration of AA10-treated abiu fruit was rel‐
atively constant (stable) over the storage period, which
may reflect reduced deterioration and senescence.
Ayon-Reyna et al. (2019) reported ascorbic acid
induced O2 consumption stabilization, inhibited pine‐
apple deterioration, and slowed senescence. From these
results, the AA10 treatment delays abiu fruit respiration
rate, thus further extends shelf life.
The ACC concentration in AA10-treated fruit tended
to be less than in the other fruit (Fig. 10a). The highest
concentration of ACC in abiu fruit occurred on day 6
for the control and AA5 treatment. In contrast, the ACC
maximum concentration for the AA10-treated fruit
occurred on day 9. The fruit senescence inhibiting
 Hort. J. Preview 7

60 AA10 AA5 control 90 AA10 AA5 control

CO2 production (mL ∙ kg-1 ∙ h-1)

a a
a
45 a
b
a a
60

ACC (mL ∙ kg -1)

b
30 a a b a b
c a a
c
b b c
a
15
30 c
a a
0 b
0 3 5 6 7 8 9 12
Days after storage 0
0 3 6 9 12
(a) Days after storage
80
O2 consumption (mL ∙ kg-1 ∙ h-1)

AA10 AA5 control (a)

a
70 a 125
a a a AA10

Ethylene production (µL ∙ kg -1 ∙ h -1)

a
60 a b AA5
a b b 100
c
a a a control
50
ab 75
a
b a
40 b b a b b
50 b b b
a a
30 c a
c b
0 3 5 6 7 8 9 12 a a
25
Days after storage

(b) 0
0 3 5 6 7 8 9 12
Fig. 9. Effect of ascorbic acid treatments on respiration; a) CO2 Days after storage
production, and b) O2 consumption of abiu fruit during storage.
Data are means of three replicates ± SE. Different letters in the (b)
same day indicate significant difference by the Duncan multiple Fig. 10. Effect of ascorbic acid treatments on; a) ACC, and b) eth‐
range test (P < 0.05). ylene production of abiu fruit during storage. Data are means
of three replicates ± SE. Different letters in the same day indi‐
cate significant difference by the Duncan multiple range test
effect of ascorbic acid is related to inhibition of ACC (P < 0.05).
production, ethylene, and fruit respiration (Martinez-
Ortiz et al., 2019). ACC production in guava coated 16
a a a
with ascorbic acid is inhibited during storage a a
a a
(Martinez-Ortiz et al., 2019). ACC synthase is a key a
b b
ethylene biosynthesis regulatory enzyme, catalyzing
SSC (°Brix)

ethylene precursor production from S-Adenosyl-

8
Methionine (SAM). Furthermore, peak ethylene pro‐
duction in the control abiu fruit occurred on day 5
(Fig. 10b). Meanwhile, peak ethylene production for the AA10 AA5 control

AA5- and AA10-treated fruit were 2 and 3 days later

than the control, respectively. Thus, the respiration rate 0
was slowed, and ethylene production decreased using 0 3 6 9 12
Days after storage
the AA10 treatment. This further supports the AA10
treatment as extending abiu fruit shelf life. Fig. 11. Effect of ascorbic acid treatments on abiu fruit SSC during
storage. Data are means of three replicates ± SE. Different
letters in the same day indicate significant difference by the
SSC and sugars Duncan multiple range test (P < 0.05).
The SSC is an important indicator determining fruit
quality and consumer acceptance (Crisosto and
Crisosto, 2005). The SSC values of the control abiu abiu fruit on day 0 was 12.54 °Brix, this value
fruit gradually increased from day 0 until day 6 of stor‐ increased 2.19, 1.68, and 1.03 °Brix by day 12 of stor‐
age, then steadily decreased until day 12 of storage age for the control, AA5-, and AA10-treated fruit,
(Fig. 11). The SSC increase in early storage is associat‐ respectively (Fig. 11). For the AA10-treated fruit,
ed with the degradation of carbohydrate polymers into exogenous ascorbic acid inhibited SSC increase, reduc‐
their sub-units. After 6 days of storage, the SSC ing the SSC by 1°Brix at the end of storage, compared
decreased due to the high metabolism in the fruit pulp to the control. The AA10 treatment efficiently delayed
and the processes leading to senescence. The SSC of starch and organic acid conversion to sugars. This
 8 S. Susanto, A. B. Arif, S. M. Widayanti and D. D. Matra

behavior was also observed in strawberries with ascor‐
bic acid treatment suppressing SSC increase by ≤ 1
°Brix compared to control strawberries (Saleem et al.,
2021).
The sucrose content in abiu fruit increased until the
climacteric peak, then decreased until the end of the
storage (Fig. 12a). This sucrose content increase is
thought to be due to starch degradation to sucrose until
the climacteric peak occurs. Starch content continues to
decrease until the climacteric peak, after which, sucrose
is hydrolyzed faster into glucose and fructose than it is
formed (Agopian et al., 2011). Exogenous ascorbic acid
treatment maintains higher sucrose content and slows
the conversion of sucrose into reducing sugars (glucose
and fructose) which stabilizes longan fruit taste and
nutritional attributes during storage (Liu et al., 2021).
10 a decreased over storage for all treatments. However,
a
a
a
b b
8 b
b c
Sucrose (%)
Exogenous ascorbic acid treatment also suppresses
polygalacturonase (PG) activity, which is responsible
for polysaccharide hydrolysis, including sucrose (Lo’ay
and El-Boray, 2018).
Fruit glucose and fructose content increased during
storage, but the glucose and fructose levels for ascorbic
acid treated fruit were lower than the control (Fig. 12b,
c). We suspect that it is caused by the reduced rate of
sucrose hydrolysis and starch breakdown. Tao et al.
(2021) reported that organic acid treatment inhibited
increases in glucose and fructose content for tomato
fruit during storage, where the acid treatment increased
sucrose phosphate synthase (SPS) enzyme activity and
decreased acid invertase (AI) enzyme activity.
TA and vitamin C
TA and vitamin C are also important markers of fruit
quality. As shown in Figure 13, TA in abiu fruit
AA10 treatment substantially inhibited TA loss during
storage. TA in the AA10-treated fruit was 3.57-fold

a higher than the control and 1.23-fold higher than the

5 c AA5 treatment on day 12 of storage. During respiration,
organic acids are converted to sugars, and their deriva‐
3 AA10 AA5 control tives or use during storage may be a contributing factor
to the TA decrease (Tilahun et al., 2019, 2020). The
0 exogenous ascorbic acid treatment of abiu fruit poten‐
0 3 6 9 12
Days after storage tially delays the consumption of organic acids inherent
(a) in the fruit, which inhibits changes in TA. Thus, the
AA10 treatment of abiu fruit maintains the TA content
4 AA10 AA5 control during the storage period.
a Vitamin C is an essential nutrient and, during storage,
3 is more susceptible to oxidation compared to other
a
Glucose (%)

a nutrients (Veltman et al., 2000). Vitamin C levels in

2 a b abiu fruit decreased during the storage period for all
b
b
b treatments (Fig. 14). However, the AA10 treatment
1 a
significantly inhibited vitamin C decreases compared
to the control and AA5 treatment. On day 12 of stor‐
0
0 3 6 9 12 age, the fruit vitamin C content was 16.15 and
Days after storage 16.33 mg·10−2·g−1 for the control and AA5 treatment,
(b)

4 AA10 AA5 control

0.20 a AA10 AA5 control
3 a
Fructose (%)

a
0.15
2 b
a a b
TA (%)

a 0.10 a
1 a b a a
a
b a
b b
0 0.05 b
0 3 6 9 12 c
Days after storage
0.00
(c) 0 3 6 9 12
Days after storage
Fig. 12. Effect of ascorbic acid treatments on; a) sucrose, b) glu‐
cose, and c) fructose levels in abiu fruit during storage. Data Fig. 13. Effect of ascorbic acid treatments on abiu fruit TA during
are means of three replicates ± SE. Different letters in the same storage. Data are means of three replicates ± SE. Different
day indicate significant difference by the Duncan multiple letters in the same day indicate significant difference by the
range test (P < 0.05). Duncan multiple range test (P < 0.05).

60
AA10 AA5
a
Vitamin C (mg ∙ 10 -2 ∙ g -1)
40 a changes in activity and protein levels of the enzymes asso‐
b a
c a
b
b a
20 c
c b
0 1–20.
0 3 6 9
Days after storage
Fig. 14. Effect of ascorbic acid treatments on abiu fruit vitamin C
levels during storage. Data are means of three replicates ± SE.
Different letters in the same day indicate significant difference
by the Duncan multiple range test (P < 0.05).
respectively. Meanwhile, for AA10-treated abiu fruit,
the vitamin C content was 1.44-fold higher than the
control. The AA10 treatment of abiu fruit preserved
vitamin C content during storage. Thus, the exogenous
ascorbic acid treatment maintains vitamin C content,
retaining fruit antioxidant properties. Arafat (2009) also
reported that, for mangoes, exogenous ascorbic acid
treatment retained vitamin C content and functioned as
an antioxidant during the storage period. Vitamin C is
an important antioxidant that can delay senescence in
fruit and reduce reactive oxygen species (Razzaq et al.,
2015).
Conclusion
In conclusion, our study demonstrated an exogenous
ascorbic acid treatment at a concentration of 10 mM
increased abiu fruit shelf life up to 12 days, nine days
longer than the control. In addition, the AA10 treatment
of abiu fruit; 1) inhibited weight loss, browning, and
MDA, 2) reduced respiration rate, ethylene production,
and phenolic oxidation, and 3) maintained vitamin C
and fruit firmness. Therefore, the AA10 treatment is an
excellent candidate as a treatment to extend abiu fruit
shelf life and maintain quality.
Acknowledgements
The author would like to thank financial support for
this research provided by Directorate General of Higher
Education, Research and Technology, Ministry of
Education, Culture, Research and Technology, Republic
of Indonesia in accordance with The Research
Implementation Assignment Agreement for the fiscal
year 2022 No.: 001/E5/PG.02.00PT/2022 and Letter of
Assignment No.: 3665/IT3.L1/PT.01.03/P/B/2022.
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