Scientia Horticulturae 188 (2015) 44–48

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Scientia Horticulturae
j o u r n a l h o m e p a g e : w w w. e l s e v i e r. c o m / l o c a t e / s c i h o r t i

Changes in fruit firmness, cell wall composition and cell wall

degrading enzymes in postharvest blueberries during storage
Hangjun Chen a , Shifeng Cao b,∗ , Xiangjun Fang a , Honglei Mu a , Hailong Yang c , Xiu Wang a
, Qingqing Xu a , Haiyan Gao a,∗∗
a
Key Laboratory of Fruits and Vegetables Postharvest and Processing Technology Research of Zhejiang Province, Zhejiang Academy of Agricultural Sciences,
Hangzhou 310021, China
b Nanjing Research Institute for Agricultural Mechanization, Ministry of Agriculture, Liuying 100, Nanjing 210014, China
c
College of Life and Environmental Science, Wenzhou University, Wenzhou 325027, China

a r t i c l e i n f o a b s t r a c t

Article history:
Blueberries are now the second most economically important soft fruit. However, they are highly per-
Received 5 January 2015
ishable and susceptible to rapid spoilage. One of the main factors limiting postharvest life of
Received in revised form 11 March 2015
Accepted 13 March
blueberries is softening. The changes of fruit firmness, cell wall degrading enzymes and cell wall
2015 composition of
Available online 3 April 2015 ‘Brilliant’ blueberry (Vaccinium ashei cv. Brilliant) were investigated in this study. The results showed
fruit firmness declined concomitantly with the increase of the content of water soluble pectin (WSP)
Keywords: during storage paralleled by a decreasing amount of sodium carbonate soluble pectin (SSP), cellulose
Blueberry and hemicellulose. Blueberries stored at low temperature (5 ◦ C) maintained higher fruit firmness
Firmness than those stored at 10 ◦ C, which was due to the lower WSP content and higher contents of SSP,
Cell wall composition cellulose and hemicellulose. Meanwhile, the lower activities of cell wall degrading enzymes such as
Cell wall degrading enzymes
polygalacturonase, cellulase, -galactosidase and -mannosidase in blueberries at 5 ◦ C were
associated with greater fruit firmness and lower WSP content as compared to those in fruit stored at
10 ◦ C.
© 2015 Elsevier B.V. All rights reserved.

1. Introduction perishable and susceptible to rapid spoilage (Cantína et al., 2012).

Blueberries, one of the most widely consumed fruit in the
world, contain high amounts of phenolic compounds, including
anthocyanins, flavonols, chlorogenic acid and procyanidins (Koca
and Karadeniz, 2009), and have been shown a wide diversity
of bioactivities such as antioxidant, antidiabetic, antimicrobial,
antiproliferative, apoptotic, liver protection, lifespan-prolonging,
anti-inflammatory, cancer preventive and cardioprotective activi-
ties (Smith et al., 2000; Faria et al., 2005; Torri et al., 2007;
Bingül et al., 2013; Bunea et al., 2013). Due to their various health
benefits, unique taste, and nutritional value, worldwide
production and con- sumption of blueberries have increased
rapidly in recent years and they have become the second most
important soft fruit species after strawberry (Giongo et al., 2013).
However, blueberries are highly
∗ Corresponding author at: Nanjing Research Institute for Agricultural Mechaniza-
tion, Ministry of Agriculture, Liuying 100, Nanjing 210014, China.
Tel.: +86 2558619521.
∗∗ Corresponding author at: Zhejiang Academy of Agricultural Sciences, Hangzhou
310021, China. Tel.: +86 57186406661.
E-mail addresses: shifengcao1@gmail.com (S. Cao), spsghy@163.com (H. Gao).
It is reported that fresh blueberries have a shelf life of 1–8 weeks
depending on stage of fruit ripeness, method of harvest, presence
of fruit disease, and storage conditions (Duan et al., 2011). One of
the main factors limiting postharvest life of blueberries is
softening (Angeletti et al., 2010), which may influence not only
the quality of the fruit, but also its storage life, transportability
and resistance of postharvest diseases (Deng et al., 2005).
Softening in any fruit is primarily due to the change in cell-
wall carbohydrate metabolism, leading to a net decrease in certain
struc- tural components (Sethu et al., 1996). During fruit
softening, the loss of firmness is associated with the decrease in
total water sol- uble pectin and the disassembly of primary cell
wall and middle lamella structures (Giongo et al., 2013).
Hemicellulose depoly- merisation and arabinose loss are the
main cell wall modifications (Vicente et al., 2007a). It is well
documented that the changes in cell wall composition and
structure results from the coordinated action of hydrolytic
enzymes in the fruit (Deng et al., 2005). Prominent enzyme,
polygalacturonase (PG), as well as a variety of glycanases and
glycosidases, plays important roles in cell wall degradation
(Sethu et al., 1996).
Several preservation technologies, including cold storage
(Connor et al., 2002), high oxygen atmospheres storage (Zheng

http://dx.doi.org/10.1016/j.scienta.2015.03.018
0304-4238/© 2015 Elsevier B.V. All rights reserved.
H. Chen et al. / Scientia Horticulturae 188 (2015) 44–48 45
46 H. Chen et al. / Scientia Horticulturae 188 (2015) 44–48

et al., 2003), allyl r m methods of Deng et al.

isothiocyanate (Wang et al., i n (2005) and Li et al. (2006).
a e Briefly, water soluble pectin
2010) and edible coating
l s (WSP) was obtained by
(Duan et al., 2011), have s
been used to maintain suspending CWM in 50 mM
bioactive compounds, reduce a sodium acetate buffer (pH
n d 6.5) for 6 h of shaking, and
deterioration, and prolong
d e collecting supernatant by
shelf life of fresh blueberries. t
However, the modification of centrifuging at 4 ◦ C. The
e
cell wall components in t water-insoluble residue was
r
r re-suspended
postharvest blueberries m
e i in 50 mM sodium acetate
during storage is still not
a n buffer (pH 6.5) containing 50
clear, and its pos- sible t a mM EDTA, shaken for 6 h,
mechanism during softening m t and centrifuged. The
is not understood. e i supernatant was collected as
Maintaining textural quality n o chelator soluble pectin (CSP).
during storage is of interest t n Sodium carbonate soluble
to the fruit growing and
pectin (SSP) was pooled by
distribution industries of Blueberry (Vaccinium Fruit firmness re-suspending the residue in
blueberries, therefore, in the ashei cv. Brilliant) fruit measurement was conducted 50 mM Na2 CO3 containing 2
present work, composition were hand- harvested from by a TA-XT plus texture
five-year-old blueberry plants mM EDTA, shaking,
modifications in the cell wall analyzer (Stable Micro
in a commercial orchard centrifuging and collecting
and changes of the activities Systems Ltd., U.K.) with a 5 the supernatant. The
of cell wall degrading located in Anji county (119◦ mm diameter stainless probe. remaining residue was re-
enzymes such as PG, 68 N, 30◦ 63 E) of Zhejiang Firmness was measured on suspended in 4 mM NaOH
cellulase, - galactosidase Province. All fruit were the equatorial region of each containing 100 mM NaBH4 ,
and -mannosidase of harvested at commercial fruit. Twenty fruit from each shaken and centrifuged. The
blueberries were measured to maturity, as deter- mined by treatment were com- pressed supernatant was collected as
explore the softening complete blue skin colour, 5 mm at a rate of 1.0 mm/s hemicellulosic fraction and
mechanism in this kind of fruit and transported within 2 h to and firmness was expressed the final residue was
during storage. the laboratory. Fruit with in kilogram per square cellulosic fraction.
uniform size and colour were centimeter (kg/cm2 ).
placed in plastic containers
2
. with snap-on lids and each
contained two hundred fruit 2.3.
Cell
(about 300 g). The containers
M wall
were divided into two groups prepa
a
t randomly. One group was ration
e then stored at 5 ◦ C and the and
r other fracti
i 10 ◦ C. Samples were taken onatio
a initially and at 7-day n
l intervals during
s s Cell wall polysaccharides
t were obtained as ethanol
a o insoluble residue using the
n r methods described by Deng
d a et al. (2005). Briefly,
g
10 g of flesh were ground,
e
m extracted by 80% (v/v) ethanol
e and main- tained in boiling
t o water to inactivate enzymes.
h f
Then the sample was
o
centrifuged after cooling and
d 4
s the residue was re-extracted
9
twice with 80% ethanol. The
2 retained residue was
d incubated overnight
.
a with 90% (v/v)
1
y dimethysulphoxide at 4 ◦ C to
.
s remove starch, and
.
then washed twice with
F
r water, chloroform–ethanol
u 2 (2:1), and ace- tone,
i . respectively. The isolated cell
t 2 wall materials (CWM) were
. dried in a vacuum oven at
m 40 ◦ C and stored over silica
a F gel in a vacuum desiccator.
t i The CWM was
e r fractionated according to the
 H. Chen et al. / Scientia Horticulturae 188 (2015) 44–48 47
The pectin content in and Sanwal (1998) with slight t
the fraction was measured modification. The reaction i
fi
by the m-hydroxydiphenyl mixture contained 2.0 ml of s
r
method (Paul and Jerome, 0.2 M sodium acetate buffer t
m
i
1982) using galac- turonic (pH 5.0), 1.0 ml of 1% (w/v) n
c
acid as standard. The cellulose solution of citrus pectin, and e
a
and hemicelluloses contents 1.0 ml of crude enzyme. The s
l
were determined using the amount of reducing sugar s
anthrone method (Vicente et released was deter- mined
a Fruit firmness is an
al., 2005) using glucose as using the dinitrosalicylate n
standard. method after reaction 30 min important quality attribute in
a
at blueberry, and excessive
l
50 ◦ C. One unit of enzyme was y softening is one of the main
2
the amount which catalyses s factors reducing quality and
.
the for- i limiting commercialization for
4
. mation of 1 g of reducing s fresh consumption (Angeletti
sugar per hour per g of et al.,
original fresh weight. All the measurements 2010). As shown in Fig. 1,
E
Cellulase activity was were conducted in triplicate. irrespective of the storage
n
determined by measuring the Data pre- sented were the temperatures, firmness of
z
y reducing sugar released from means ± SD values. All blueberries increased during
m statistical analyses were the first 7 days of storage, and
carboxymethyl cellulose
e performed with using SAS declined gradually
(Deng et al., 2005). The
statistical software 8.01 (SAS afterwards. Fruit stored at 5 ◦
reaction mixture contained
institute, Cary, NC). C maintained greater
e 2.0 ml of 1% (w/v) solution of
x firmness than those at 10 ◦ C
car- boxymethyl cellulose
t after 7 days of storage.
and 0.5 ml of crude enzyme. 3
r
The amount of reducing sugar .
a 3
released was determined
c .
t using the dinitrosalicylate R 2
i method after reaction 30 min e .
o at 50 ◦ C. One unit of enzyme s
n was the u
amount which catalyses the l C
formation of 1 mg of t h
a a
reducing sugar per hour per g s
n n
d of original fresh weight. g
-Galactosidase and a e
-mannosidase activity was n s
a
determined by measuring d
s
s p-nitrophenol released i
a from p-nitrophenyl- - d n
y galactopyranoside and p- i
nitrophenyl- s
c c
Two grams of fruit tissues -mannopyranoside,
u e
were treated with liquid respectively (Sethu et al., l
s
nitrogen, pulverized and 1996; Deng et al., 2005). The s l
extracted in 10 mL of 0.2 M reaction mixture contained i
sodium acetate buffer (pH 5.0) 0.5 ml of sodium acetate o w
containing 1 mM EDTA-Na, 5% buffer (0.2 M, pH 5.0), n a
polyvinylpyrrolidone (w/v). 0.18 ml of 16 mM p- l
The enzyme extract was nitrophenyl- 3 l
obtained by centrifugation at -galactopyranoside and 0.12 .
10,000×g for ml of crude enzyme. The 1
c
2 amount of p-nitrophenol was .
o
0 measured after reaction 90 m
min at 37 ◦ C. One unit of C p
m h o
i enzyme was the amount
a s
n which catalyses the formation
n i
of 1 mol of p-nitrophenol per g t
a hour per g of original fresh e i
t weight. s o
n
4
2 i
◦ . n Previous work analyzing
5 cell wall changes in blueberry
. fruit during development
C f
. r showed that pectin
PG activity was assayed by S u solubilization increased
the method described by t i during ripening (Vicente et
Deng et al. (2005) and Pathak a t al., 2007b). Less attention has
48 H. Chen et al. / Scientia Horticulturae 188 (2015) 44–48
been
H. Chen et al. / Scientia Horticulturae 188 (2015) 44–48 49
50 H. Chen et al. / Scientia Horticulturae 188 (2015) 44–48

2.5 three pectin-rich fractions as WSP, CSP, and SSP and hemicellulose
and cellulose. Among them, WSP was thought to represent wall
o
5 C polymers solubilized in vivo but remaining in the apoplast which
2.0 10 oC could be obtained by water extraction of the purified cell wall,
whereas CSP and SSP are generally considered to be enriched for
* * * ionically and covalently bound pectins, respectively. Typically, dur-
ing fruit softening, depolymerisation of cell wall polysaccharides
Firmness (N)

1.5
1.0
0.5
0.0
0 7
Fig. 1. Changes on firmness in blueberry fruit during storage at 10 ◦ C and 5 ◦ C. Val-
ues are the means ± SE. Vertical bars represent the standard errors of the means.
Asterisks indicate significant differences between the fruit stored at 10 ◦ C and 5
C (Duncan’s multiple range test, * P < 0.05).
paid to the changes in pectin, hemicellulose and cellulose con-
tent of blueberries after harvest. In this study, we examined the
possible role that cell wall modification might play in response
to fruit softening during storage. Cell walls were extracted to give
* was accompanied by increases in the levels of WSP, whilst the lev-
*
els of CSP, SSP, hemicellulose and cellulose decreased (Carrington
* et al., 1993; Vicente et al., 2007a). In our present study, the pectin
fractions changed significantly in blueberries during storage at 5 ◦
C and 10 ◦ C. The amount of WSP increased gradually with
storage time paralleled by a decreasing amount of SSP (Fig. 2A and
B). Mean- while, the amount of CSP increased during the first 21
days, and then decreased afterwards (Fig. 2C). These changes
suggested that, similar to the other fruit, pectins, as indicated by
uronic acid con- tent, were solubilized from the wall during
softening in blueberries, indicating the possible alterations in the
14 21 28 35 42 49 cross-linking of carbohy- drates in the cell walls (Manning,
1993). Moreover, it should be noted that the increase in fruit
Storage time firmness in blueberries during first
(day) 7 days of storage observed in our present study might be resulted
from the CSP accumulation (Fig. 2C) in this fruit regardless of the
storage temperatures.
◦ Additionally, in our present study, we also observed that blue-
berries stored at 5 ◦ C experienced higher levels of SSP and CSP
but lower WSP content as compared with those at 10 ◦ C, which
were 31.2% and 116.7% higher in levels of SSP and CSP, respec-
tively, and 14.6% lower in WSP content than those in fruit stored
at 10 ◦ C at the end of storage. Thus, results here suggested that
low temperature storage somehow prevented solubilization of
these

0.6 2.8
A * * * B

Na 2CO 3soluble pectin (%)

Chelator soluble pectin (%) Water soluble pectin (%)

* * * * * 2.1
0.4
* *
1.4
o
0.2 o 5 C
5 C
o
o
10 C 0.7
10 C

0.0 0.0
1.8 1.5
C D
Hemicellulose (%)

* *
1.2 * 1.0
* * * *
0.6 * 0.5

0.0 0.0

3.5 E
3.0
*
*
Cellulose (%)

*
2.5 *
2.0

1.5

1.0
0 7 14 21 28 35 42 49
Storage time
(day)

Fig. 2. Changes in water-soluble pectin (A), sodium carbonate-soluble pectin (B), chelator soluble pectin (C), hemicellulose (D) and cellulose (E) contents of blueberry fruit
during storage at 10 ◦ C and 5 ◦ C. Values are the means ± SE. Vertical bars represent the standard errors of the means. Asterisks indicate significant differences between
 H. Chen et al. / Scientia Horticulturae 188 (2015) 44–48 51
the fruit stored at 10 ◦ C and 5 ◦ C (Duncan’s multiple range test, * P < 0.05).
 10000 15000
A * * B
8000 * 12000

Polygalacturonase

(mg h-1g-1 FW)

* * * * *

(μg h-1g-1 FW)

* *

Cellulase
6000 9000
* * *
4000 6000

2000 o
5 C
o
5 C 3000
o o
10 C 10 C
0 0
800 1000
C
* D
* 800
* * *

(μmol h g FW)
600 *

α-Mannosidase
*
(μmol h g FW)
β-Galactosidase

* * * 600

-1
-1
-1 -1

400
400
200
200

0 0
Storage time
(day)

Fig. 3. Changes in activities of polygalacturonase (A), cellulase (B), -galactosidase (C) and -mannosidase (D) of blueberry fruit during storage at 10 ◦ C and 5 ◦ C. Values
are the means ± SE. Vertical bars represent the standard errors of the means. Asterisks indicate significant differences between the fruit stored at 10 ◦ C and 5 ◦ C (Duncan’s
multiple range test, * P < 0.05).

polymers. It was reported that the architectural strength of cell
wall can be increased by the cross-linking of pectin and the
associ- ation between pectin and extensin proteins (Vicente et al.,
2007a). As higher amount of ionically and covalently bound
pectins (CSP and SSP) was maintained in blueberries stored at 5 ◦
C, it probably cross-linked to other cell wall polymers and
strengthened the cell wall, which was associated with the greater
firmness.
Increasing evidence highlighted solubilization and
depolymer- ization of hemicellulose and cellulose also played
important roles in fruit softening (Wakabayashi et al., 2000).
Vicente et al. (2007b) demonstrated hemicellulose degradation
was among the main modifications of cell wall disassembly in
blueberry fruit dur- ing development. Results found in the
present work show that contents of hemicellulose and cellulose
decreased gradually in blueberries during storage regardless of
the storage tempera- tures, lower temperatures maintained
higher levels of these two components in blueberries, which
were 50.0% and 66.7% higher, respectively, than those in fruit at
10 ◦ C after 49 days of storage (Fig. 2D and E). It was
documented that the rigidity of cellular walls was probably
attributed to the cellulose-hemicellulose net- work structure,
formed by hydrogen-bond and crosslinks between cellulose
microfibrils (Wakabayashi et al., 2000). The breakdown of
hemicellulose was revealed to result in the disassembly of the
cellulose-hemicellulose network and fruit softening (Cheng et al.,
2009). On the other hand, due to the crystalline nature of
cellulose, its change was related to the resistance of cell wall to
enzymatic degradation (Zhou et al., 2011). Taken together, our
results suggest that the continuous degradation of hemicellulose
and cellulose is of importance to fruit softening of blueberries
during storage as well as an array of other fruit (Wakabayashi et al.,
2000; Zhou et al., 2011; Wang et al., 2015). Cold storage
suppressed the depolymerization and degradation of cellulose
and hemicellulose, thus delaying fruit softening.
3.3. Changes in cell wall degrading enzymes
It was increasingly apparent that softening and textural changes
were brought about by the actions of a multitude of cell
wall-localized enzymes acting on specific, potentially highly local-
ized substrates (Brummell et al., 2004). PG hydrolyzes pectin acid
along with the main chain of polygalacturonic acid, causing pectin
degradation, cell wall dissolution, and ultimately, fruit softening
(Wei et al., 2010). Cellulase degrades both cellulose and -1,4-
glucan backbone of xyloglucan, a hemicellulosic polysaccharide
abundant in cell walls of dicotyledons (Vicente et al., 2007a; Bu
et al., 2013). -Galactosidase has been characterized in association
with the removal of galactosyl residues from cell wall polymers
during fruit softening (Wei et al., 2010). The removal of pec-
tic galactan side-chains was an important factor in the cell wall
changes leading to ripening-related firmness loss (Payasi et al.,
2009). Glycosidases such as -mannosidase and -galactosidase
are a class of carbohydrate hydrolyses that act on short chain
oligosaccharides, which may be present in glycoproteins, and on
carbohydrate heteromers or homomers (Vicente et al., 2007a). In
order to reveal the mechanism involved in fruit softening of blue-
berries during storage, the activities of four cell wall degrading
enzymes such as PG, cellulase, -galactosidase and
-mannosidase were measured. Regardless of storage
temperatures, activities of all these four enzymes in blueberries
showed similar changes dur- ing storage, which increased and
peaked firstly and then declined
afterwards (Fig. 3). Low temperature at 5 ◦ C remarkably
suppressed
activities of the four enzymes and maintained lower WSP con-
tent and higher contents of hemicellulose and cellulose
compared with the fruit stored at 10 ◦ C, which suggested that
cold stor- age inhibited cell wall degradation by inhibiting its
degrading enzymes. At the end of the storage, activities of
cellulase, - galactosidase and -mannosidase in fruit at 5 ◦ C
were 3.2-, 1.5- and 1.2-fold, higher than those at 10 ◦ C,
respectively. Vicente et al. (2005) also found that softening
process was delayed via the inhibition of cell wall degrading
enzymes including PG and - galactosidase in heat treated
strawberries, which confirmed that cell wall degrading enzymes
play an important role in fruit soften- ing. Therefore, our results
indicated that the effect of cold storage on delaying softening in
blueberries fruit may be partially due to relatively lower
activities of PG, cellulase, -galactosidase and - mannosidase.
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